FOUR JUMPING SPIDER GENERA OF THE COCALODES-GROVV ARE MONOPHYLETIC WITH GENERA OF THE SPARTAEINAE (ARANEAE: SALTICIDAE)

A.C RODRIGO1 & R.R. JACKSON2

department of Zoology, School of Biological Sciences, University of Auckland, Private Bag 92019, Auckland, New Zealand & 2Department of Zoology, University of Canterbury, Private Bag, Christchurch, New Zealand.

(Received 6 January 1992; revised and accepted 30 March 1992)

ABSTRACT

Rodrigo, A.G. & Jackson, R.R. (1992). Four jumping spider genera of the Cocalodes-group are monophyletic with genera of the Spartaeinae (Araneae: Salticidae). New Zealand Natural Sciences 19: 61-67.

Wanless (1984a) erected a new sub-family, the Spartaeinae, to contain all members of the incorrectly named Boethinae except the genera Allococalodes, Cocalodes, Holcolaetis, and Sonoita. These were excluded from the sub-family because they possessed a median tegular apophysis on the male palp, a feature which Wanless believed to be synapomorphy of the group, and one which separated the group from members of the Spartaeinae. Our phylogenetic analysis of the extant members of the Spartaeinae, and the "Cocalodes-group" indicate that the latter are monophyletic with other genera of the sub-family. An amended definition of the sub-family is provided.

KEYWORDS: Spartaeinae - "Cocalodes-group" - phylogeny - monophyly - quasi-polymorphic coding.

INTRODUCTION however, that these genera are closely affiliated with members of the Spartaeinae and another Wanless (1984a) erected the jumping spider allied sub-family, the Lyssomaninae (Wanless sub-family Spartaeinae to accomodate members 1984a). of the incorrectly named Boethinae. He also These three groups are considered to be removed the genera Cocalodes Pocock, Sonoita primitive salticids (Blest & Carter 1987), and our Peckham & Peckham, and Holcolaetis Simon interest in them stems from a desire to test the from the sub-family on the grounds that males of hypothesis first proposed by Jackson & Blest these genera possess a median apophysis on the (1982) that jumping spiders evolved from web- palpal tegulum, a feature not possessed by other building ancestors. As a consequence, we recon­ genera of the Spartaeinae. Wanless (1984a) sug­ structed the phylogeny of the Spartaeinae using gested that another sub-family should be erected morphological characters, and we included the to contain these aberrant genera and a closely related genus, Allococalodes Wanless, and he in­ genera Cocalodes, Allococalodes, Holcolaetis, and formally referred to them collectively as the "Co- Sonoita in the analysis. Our results indicate that calodes-group". He was later to reassess this the "Cocalodes-gpoxxip" forms a monophyletic claim on the grounds that Holcolaetis and Sonoita clade within the Spartaeinae and, in accordance share little in common with Cocalodes and Alloco­ with current phylogenetic systematic practice, calodes, apart from the presence of a tegular should be included as members of the sub-family. median apophysis, the homology of which he Here, we describe the phylogenetic analysis, as doubted (Wanless 1985). There is little doubt, well as the implications of its results. 62 New Zealand Natural Sciences 19 (1992)

METHODS AND MATERIALS or more groups, with at least two groups having more than one taxon as a member, were used. TAXA STUDIED Characters which satisfy this criterion are phylo­ Descriptions ofthe taxa listed in Table 1 were genetically informative (sensu Hennig 1965). obtained from published literature (cited in Table Characters which are monomorphic for all taxa 1). A list of characters were obtained from these offer no information on within-group relation­ descriptions and these are described below. ships. Characters for which there is only one partition with more than one taxon are similarly MORPHOLOGICAL CHARACTERS uninformative, for it implies that all other taxa The following morphological characters were possess unique character states (ie. autapomor- used to construct a taxon-character dataset. The phies). reader is directed to the excellent papers by Wanless (1984a, 1984b) for more complete de­ 1. Posterior median eyes (PMEs): 1-small and scriptions ofthe characters. The method of maxi­ vestigial; O-large (Wanless 1984a, p. 137). mum parsimony (Kluge & Farris 1969, Wiley Only some extant and extinct (ie. amber fossil) 1981) was used to reconstruct the phylogeny ofthe members of the Spartaeinae, Lyssomaninae group. As is appropriate for this method, only and Cocalodes-group possess large and func­ characters whose states partition the taxa into two tional PMEs. These are vestigial in other advanced salticids. Wanless (1984a) notes that with the exception of Taraxella, all salti­ Table 1. List of genera studied. Asemonea is a member ofthe cids can be placed easily into one or the other Lyssomaninae, and was included as an outgroup. Allococal- category. odes, Cocalodes, Holcolaetis, and Sonoita have been infor­ 2. PME in relation to anterior median/anterior mally referred to by Wanless as the "Cocalodes-group". lateral eyes (AMEs/ALEs) and posterior lat­ eral eyes (PLEs): 1-closer to AME/ALEs GENUS REFERENCE than to PLEs; 0-closer to PLE (Wanless 1980, p. 217). The closeness of the PMEs to the ALLOCOCALODES Wanless Wanless 1982 anterior eyes is most marked in the lyssoman­ ASEMONEA Wanless 1980 ine genera Asemonea O.P.-Cambridge, Go- BRETTUS Thorell Wanless 1979 leba Wanless, Macopaeus Simon, and Pan­ COCALODES Pocock Wanless 1982 disus Simon, where the tubercles on which the COCALUS Koch Wanless 1981a PMEs are situated abut those which hold the CYRBA Simon Wanless 1984b ALEs. The PMEs of some spartaeine genera GELOTIA Thorell Wanless 1984a are also found close to the ALEs, although not Wijesinghe, 1991b to the extent found in the lyssomanine genera. HOLCOLAETIS Simon Wanless 1985 3. Femoral organ: 1-present; 0-absent (Wanless MELEON Wanless Wanless 1984a 1984a p. 139). This structure takes the form of MINTONIA Wanless Wanless 1984a a perforated region situated either on a tu­ NEOBRETTUS Wanless Wanless 1984a bercle or in a gully on the underside of the PHAECIUS Simon Wanless 1981b femora of Legs I in the males of some genera. Wijesinghe 1991a An amorphous exudate is often associated PORTIA Karsch Wanless 1978 with the organ, and it may have a pheromone Wanless 1984a secreting function. SONOITA Peckham & Peckham Wanless 1985 4. Minute tarsal and metatarsal setae: 1-present; SPARTAEUS Thorell Wanless 1984a 0-absent (Wanless 1984a, p. 169; 1984b, p. Wanless 1987 450). Numerous minute setae may be found TARAXELLA Wanless Wanless 1984a on the underside of tarsi and metatarsi of Wanless, 1987 some genera. VEISSELLA Wanless Wanless 1984a 5. Retrolateral tibial apophysis (RTA): 1-mem- YA GINUMANIS Wanless Wanless 1984a braneous base present; 0-membraneous base A.G. Rodrigo & R.R. Jackson: Phylogeny of jumping spiders of Cocalodes-group 63

absent (Wanless 1984a, p. 139). The RTA is presence of structures (30-40 um in length) found on the male palpal tibia. The presence which resemble mussel shell valves. These of a membraneous base to the RTA allows it fields are situated on the dorsal surface ofthe to have some mobility. This feature is only abdomen, and are associated with pores of present in some genera. some form or other, suggesting a secretory 6. Ml apophysis of the distal haematodocha: 0- function. They are difficult to find in all but absent, 1-minute lobe, 2-large and petal-like, fresh specimens, and have only been found in 3-long and filamentous, 4-membraneous some species. We have been conservative in patch (Wanless 1984a, p. 140). The distal our estimation of their presence, and have haematodocha of the male palps of some only recorded them as present in those genera genera are made up of two apophyses which with species which are known to possess these are labelled by Wanless (1984a) as Ml and fields. All other genera have been treated as M2. The reader is referred to Wanless not possessing these fields. (1984a) for more complete descriptions of 12. Cymbial flange: 0-absent; 1-present (Wanless these structures, but it is noteworthy that these 1978, p. 84). A flange-like protuberance is apophyses are absent from the Cocalodes- found on the pro- or retrolateral dorsal sur­ group. Wanless (1984a) therefore suggested face at the base of the cymbium of some that the presence of Ml and M2 may be a genera. synapomorphy (ie. a shared-derived charac­ 13-16. Number of pro-marginal teeth on chelic­ ter) of the Spartaeinae. erae (Wanless 1984a, p. 138). Salticids, like 7. Median tegular apophysis: 1-present; 0-ab­ many other spiders, possess dentate armature sent (Wanless 1982, p. 264; Wanless 1984a, p. on their chelicerae. The numbers of these 141). This apophysis takes the form of a bifid cheliceral teeth have often been used as di­ prong arising from the palpal tegulum, and is agnostic characters. However, because ofthe only found in members of the "Cocalodes- variability in numbers of these teeth between group", as well as some lyssomanine genera. species within genera, such characters have 8. Tegular furrow: 0-lacking; l-short and shal­ recently been rejected as having systematic low; 2-deep (Wanless 1984a, p. 140). Part of value. Nonetheless, while the numbers of the tegulum, usually found on its dorsal side. cheliceral teeth may be variable, this variabil­ It is often obscured by other structures of the ity, as measured by its range, may itself be palp, and information on this feature is not phylogenetically informative. always available. Wanless (1984a) makes the point that the furrow is found on the palps of We have coded these continuous characters all male spartaeines, and considers it to be a using Quasi-Polymorphic Coding (QPC; Rodrigo synapomorphy of the sub-family. & Ritchie in prep.), a coding method which takes 9. Palpal patellar protuberance: 0-lacking; 1- account of the range of a character and its overlap membraneous; 2-rigid; ?-variable (Wanless amongst taxa. Briefly, for any continuous charac­ 1984a, p. 141). The patellar protuberances, ter, QPC treats the regions of overlap between two together with tibial protuberances, are part of or more taxa, and the regions unique to each taxon the architecture of the male palps and define their potential for articulation. as separate quasi-alleles. For any given area of 10. Eyes on well-developed tubercles: 0-yes; l-no. overlap, all taxa which share that region are coded Many spartaeine and lyssomanine genera as having the same quasi-allele whereas at the have their ALEs, PMEs, and PLEs set on well unique areas only one taxon possesses the quasi- developed tubercles. In others, however, allele and this is taken to be uninformative. these tubercles are poorly developed or lack­ Quasi-alleles are coded as being absent from a ing. taxon (character state 0) or present in the taxon 11. Mytiliform fields: 0-absent; 1-present (character state 1). Fig. Ia illustrates how the (Wanless 1984b, p. 446). These "mytiliform" method is used to code the numbers of pro-mar­ fields are so-called by Wanless because of the ginal teeth. 64 New Zealand Natural Sciences 19 (1992)

17-23. Number of retromarginal teeth on chelic­ acter. In order to obtain a directed evolutionary erae (Wanless 1984a, p. 138). Again, this hypothesis, the trees were rooted using the out­ character has been coded using QPC (Fig. Ib). group method (Watrous & Wheeler 1981). Ac­ cording to this method, a closely related sister PHYLOGENETIC ANALYSIS taxon (= the outgroup) of the group under study A taxon-character dataset was constructed by (= the ingroup) is selected. For each character, if assigning to each taxon, a character state for every a state is found in both the ingroup and the out­ character (Table 2). Where information was group then that state is treated as ancestral (= unavailable on any particular character for a given plesiomorphy). Other states of the character are taxon, that character was coded as "unknown" or taken to be derived ( = apomorphies). If two taxa "missing" (this is indicated by a "?" in Table 2). share derived states, it is an indication of phylogen­ The computer program PAUP 3.0L (Phylo­ etic relationship. Outgroup analysis is imple­ genetic Analysis Using Parsimony: Swofford mented automatically in PAUP 3.0L. We chose 1990) was used to reconstruct the phylogeny ofthe the genusAsemonea, a member ofthe Lyssoman­ group under the principle of maximum-parsi­ inae, as an outgroup. mony. According to the parsimony criterion, a Eleven most parsimonius trees were found tree is selected as a plausible phylogenetic hy­ using the tree-bisection reconnection (TBR) op­ pothesis if it requires the fewest number of charac­ tion in PAUP 3.0L (Swofford 1990). All trees had ter state changes (Kluge & Farris 1969, Farris a length (ie. number of character changes) of 64 1983). All characters were treated as unordered, and a consistency index of 0.422. The consistency ie. no prior assumption was made about the evolu­ index hes in the range (0,1) and can be taken as a tionary sequence of transformations of each char­ measure of how well the characters are indicators

Table 2. Morphological character-taxon dataset. See text for details of the characters and their codes. Missing or variable information is indicated by "?".

Characters

11111111112222 Taxon 12345678901234567890123

Brettus 0 1 1 1 0 2 0 ? 2 0 0 9 1 0 0 0 1 1 0 0 0 0 0 Cocalus 0 1 0 1 0 2 0 1 0 0 1 0 1 0 0 0 0 1 1 0 0 0 0 Cyrba 1 0 0 1 0 1 0 9 ? 9 1 1 1 0 0 0 1 1 1 0 0 0 0 Gelotia 1 0 1 1 0 0 0 9 2 0 1 0 1 0 0 0 0 1 1 0 0 0 0 Meleon 0 0 0 1 1 3 0 2 ? 0 0 0 1 0 0 0 1 1 1 1 0 0 0 Mintonia 0 1 1 0 0 3 0 1 0 0 1 0 1 0 0 0 0 0 1 1 1 0 0 Neobrettus 0 1 0 0 0 3 0 ? 2 0 0 0 1 0 0 0 0 1 1 0 0 0 0 Phaecius 0 0 0 1 1 3 0 1 2 1 1 1 1 0 0 0 1 1 1 0 0 0 0 Portia 0 0 0 1 9 3 0 1 2 1 1 1 1 0 0 0 0 0 0 1 0 0 0 Spartaeus 0 1 1 1 0 3 0 ? 0 0 0 0 0 0 1 1 0 0 0 0 1 1 1 Taraxella ? 1 0 0 0 1 0 2 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 Veisella 0 0 0 1 0 3 0 1 0 0 0 0 1 0 0 0 1 1 0 0 0 0 0 Yaginumanis 0 1 0 0 0 3 0 ? 0 0 0 0 1 0 0 0 0 0 1 1 0 0 0 Cocalodes 0 1 0 0 0 4 1 9 0 0 0 1 1 0 0 0 1 0 0 0 0 0 0 Allococalodes 0 1 0 0 0 4 1 9 0 0 0 1 0 1 0 0 0 0 0 0 0 0 0 Holcolaetis 1 1 0 0 0 4 1 9 9 1 1 9 0 1 1 1 0 0 0 1 1 1 1 Sonoita 0 1 0 0 0 4 1 9 9 1 1 1 1 1 1 0 0 0 1 1 0 0 0 Asemonea 0 1 0 0 0 ? 0 9 0 0 0 0 1 0 0 0 1 0 1 1 1 0 0 A.C Rodrigo & R.R. Jackson: Phylogeny of jumping spiders of Cocalodes-group 65

Quasi-aileles present in taxon , Quasi-alleles u present in taxon 2i3 :4 :5 :6 :7 : 2J3;4: 5; 6: 7;8;9:10:11: Allococalodes Allococalodes !—! :::::;: Asemonea Asemonea ; j—j" j—]—|11H" ''•' *• *."•" ": Brettus Brettus ' ; j—•—j : : : : ; ' : Cocalodes Cocalodes i i— : : : : : : : •' Cocalus Cocalus ' : I—i—J .--;--.--.---.-•-; Cyrba Cyrba ; i—i—;—i : : ; : ; Gelotia Gelotia • ; j—-j—f ; • :**:""*;"*: Holcolaeiis Holcolaetis •'• ','.'. i—i—;—•—;—i '• Meleon Meleon : t—i—:—:—j : ; : : > Mintonia Mintonia • •" • \—[—j—I ' ']"•': Neobrettus Neobrettus • ; j—\—j • • * :" >'' *|" * ; Phaecius Phaecius : )—:—:—j : : ; : : : i..,-..'..i..'...'..i.....'...•... • Portia Portia : : : : \—j : : : : \ Spartaeus I ; ! Spartaeus : : : : : i—:—r-:—r-i Sonoita Sonoita • •"" | j "\ "j""'' V T "f"" - taraxella taraxella :::::: i—•—f* "":*"' Veissella Veissella : l ":" I :::;:": j Yaginumanis Yaginumanis \ : .—:—! : ; : : '

Figure 1. Quasi-polymorphic coding: a) numbers of pro-marginal cheliceral teeth; b) numbers of retro-marginal cheliceral teeth. Horizontal bars represent quasi-alleles present in each taxon. Quasi-alleles are not necessarily related in a one-to-one manner with the character states. For instance, in (b), some taxa have members with both 8 or 9 cheliceral teeth. However, there are no taxa in which members have 8 teeth without other members having 9, or vice versa. Therefore, "8 or 9 cheliceral teeth" is represented by one quasi-allele.

of phylogenetic relationship. A consistency index workers at the Australian National University, of 1 indicates that every character is a perfect indi­ Canberra, have studied the retinal organisation of cator of phylogenetic affiliation. A strict consen­ the principal eyes of a number of jumping spiders, sus of the ll trees, showing only those clades sup­ including spartaeines and members of the genus ported by all most parsimonius trees, indicate that Allococalodes, Their studies indicate that the Cocalodes, Allococalodes, Holcolaetis, and retinal organisation found mAllococalodes is very Sonoita are part of the Spartaeinae and form a similar to that of other spartaeines. monophyletic group with two spartaeine genera, Given the support that a phylogenetic analysis Spartaeus and Taraxella (Fig. 2). and ultrastructural information offer for the monophyly ofthe spartaeines and the "Cocalodes- DISCUSSION group", there is no compelling reason not to in­ clude the latter as members of the Spartaeinae. The monophyly ofthe "Cocalodes-group" and Diagnostically, as well, the two groups are very the Spartaeinae is supported not only by our phy­ similar: for instance,Wanless (1984a), in his diag­ logenetic results, but also by studies on eye ultras­ nosis ofthe Spartaeinae, noted that to separate the tructure (Blest & Carter 1987, Blest et ai 1990, A. females Cocalodes-group from those ofthe Spar­ D. Blest pers. comm.). David Blest and co­ taeinae, literature descriptions, particularly ofthe 66 New Zealand Natural Sciences 19 (1992)

/ Brettus Wanless, Veissella Wanless, Yaginumanis /s Cocalus Wanless. Extinct: Almolinus Petrunkevitch, Cenattus Z/^- Cyrba Petrunkevitch, Eolinus Petrunkevitch, Paral- /^y^^\^ Phaecius inus Petrunkevitch, Prolinus Petrunkevitch. /C^l^ ^ Portia ACKNOWLEDGEMENTS / ^V\\7^" Gelotia We thank F. R. Wanless for examining and / xX^ Meleon commenting on an earlier morphological dataset y{ X/^ Neobrettus which included Cocalodes but not Allococalodes, /\\\ Veissella Holcolaetis, and Sonoita. Thanks also go to Don Cowley and Ian Henderson who offered helpful \Z\^\\ / Spartaeus comments on the manuscript, and L. Dunn who \^\\\ /^ Taraxella assisted with its preparation.

\\s\\>/ y Cocalodes REFERENCES \. \J\\// Allococalodes Blest, A.D. & Carter, M. (1987). Morphogenesis \\\\f/ Holcolaetis of a tiered principal retina and the evolution of \ \ \. Sonoita jumping spiders. Nature 328:152-155. \ \• Yaginumanis Blest, A.D., O'Carroll, D.C. & Carter, M. (1990). Comparative ultrastructure of Layer I recep­ \\ Mintonia tor mosica in principle eyes of jumping spi­ ^ Asemonea ders: the evolution of regular arrays of light guides. Cell and Tissue Research 262:445-460. Figure 2. A strict consensus of the ll most-parsimonious phy­ Farris, J.S. (1983). The logical basis of phylogen­ logenetic trees of the Spartaeinae and "Cocalodes-group". etic Systematics. In Advances in Cladistics, Vol. 2, Proceedings ofthe Second Meeting of the epigynes, need to be consulted. Furthermore, Willi Hennig Society (eds. N.l. Platnick & VA. separating the Spartaeinae and "Cocalodes- Funk). Columbia University Press, New group" from the Lyssomaninae is not difficult: York. members of the latter sub-family have their eyes Kluge, A. & Farris, J.S. (1969). Quantitative arranged in four rows, and have a typically lyso- phyletics and the anurans. Systematic Zoology maniform carapace (see Wanless 1980 for details). 18: 1-32. The following, then, is an amended classification Jackson, R.R. & Blest, A.D. (1982). The biology of the sub-family Spartaeinae. of Portia fimbriata, a web-building jumping spider (Araneae: Salticidae) from Queens­ Sub-family SPARTAEINAE Wanless, 1984 land: utilisation of webs and predatory Amended definition. As defined by Wanless versatility. Journal of Zoology London 196: (1984) but with the following amendments: 255-293. Body lengthy 3.0 mm to 16.0 mm; Swofford, D.L. (1990). PAUP: Phylogenetic Chelicerae, retromargin with two to ten teeth; Analysis Using Parsimony, Ver. 3.0L. Com­ Type genus, Spartaeus Thorell; puter program and manual. Illinois Natural Other genero, History Survey, Illinois. Extant: Allococalodes Wanless, BrettusThorcll, Wanless, F.R. (1978). A revision of the spider Cocalodes Pocock, Cocalus Koch, Cyrba genus Portia (Araneae: Salticidae). Bulletin of Simon, Gelotia Thorell, Holcolaetis Simon, the British Museum of Natural History (Zool­ Meleon Wanless, Mintonia Wanless, Neobret­ ogy) 38: 219-223. tus Wanless, Phaecius Simon, Portia Karsch, Wanless, F.R. (1979). A revision of the spider Sonoita Peckham & Peckham, Taraxella genus Brettus (Araneae: Salticidae). Bulletin A.C Rodrigo & R.R. Jackson: Phylogeny of jumping spiders of Cocalodes-gpoup 61

ofthe British Museum of Natural History (Zo­ genus Cyrba (Araneae: Salticidae), with the ology) 35: 127-200. description of a new presumptive pheromone Wanless, F.R. (1980). A revision of the spider dispersing organ. Bulletin ofthe British Mu­ genera Asemonea and Pandisus (Araneae: seum of Natural History (Zoology) 47: 445- Salticidae). Bulletin ofthe British Museum of 481. Natural History (Zoology) 39: 213-257. Wanless, F.R. (1985). A revision of the spider Wanless, F.R. (1981a). A revision of the spider genera Holcolaetis and Sonoita (Araneae: genus Cocalus (Araneae: Salticidae). Bulletin Salticidae). Bulletin ofthe British Museum of ofthe British Museum of Natural History (Zo­ Natural History (Zoology) 48:249-278. ology) 41:253-261. Wanless, F.R. (1987). Notes on spiders of the Wanless, F.R. (1981b). A revision of the spider family Salticidae. 1. The genera Spartaeus, genus Phaecius (Araneae: Salticidae). Bulle­ Mintonia, and Taraxella. Bulletin of the British tin of the British Museum of Natural History Museum of Natural History (Zoology) 52:107- (Zoology) 41: 199-212. 137. W7anless, F.R. (1982). A revision of the spider Watrous L.E. & Wheeler, Q.D. (1981). The out­ genus Cocalodes with a description of a new group comparison method of character related genus (Araneae: Salticidae). Bulletin analysis. Systematic Zoology 30:1-11. ofthe British Museum of Natural History (Zo­ Wijesinghe, D.P. (1991a). New species of ology) 42:263-298. Phaecius from Sri Lanka, Sumatra and the Wanless, F.R. (1984a). A revision of the spider Phillippines (Araneae: Salticidae). Bulletin of sub-family Spartaeinae nom. n. (Araneae: the British Arachnological Society 8: 249-255. Salticidae) with descriptions of six new Wiley, E.O. (1981). Phylogenetics: the theory and genera. Bulletin of the British Museum of practice of phylogenetic Systematics. Wiley & Natural History (Zoology) 46: 135-205. Sons, New York. Wanless, F.R. (1984b). A revision of the spider