ANNALES ZOOLOGICI (Warszawa), 2002, 52(3): 459-464

ZOOGEOGRAPHY OF SALTICIDAE (ARACHNIDA: ARANEAE) OF – FIRST APPROACH

MAREK ¯ABKA1, SIMON D. POLLARD2 and MATHEW ANSTEY3

1Katedra Zoologii Akademii Podlaskiej, 08-110 Siedlce, Poland, e-mail: [email protected] 2Canterbury Museum, Rolleston Avenue, 8001 Chrischurch, New Zealand, e-mail: [email protected] 3Department of Zoology, University of Canterbury, Private Bag 4800, Christchurch, New Zealand, e-mail: [email protected]

Abstract.— According to our unpublished data some 30 genera and 200 species of Salticidae can be expected in New Zealand. The fauna is highly endemic, both on a generic and a specific levels. The most diverse are two groups of genera: Trite minax [=planiceps] and “Trite” auricoma are the best known representatives of every group. The relationships between Salticidae of New Zealand and , are limited to single representatives of Opisthoncus, Holoplatys, , Helpis, “Lycidas”, “Clynotis” and Hypoblemum. Wide-spread genera are represented by Neon and Bianor and pantropical Hasarius adansoni is found in the warmer climate of North Island. To a limited extent New Zealand is a source of fauna for other Pacific archipelagos, for example species of Trite are found in and . Ë Key words.— Biogeography, Salticidae, New Zealand.

INTRODUCTION MATERIALS AND METHODS

Of about 50 salticid species formally recorded in New The material we analysed covered various habitats Zealand (¯abka and Anstey in press), great majority throughout New Zealand, including the coastline, alpine were described in the nineteenth century, only some areas, subantarctic islands, Nothofagus and subtropical have been properly diagnosed and are recognisable. forests, and was provided by the following New Zealand Most species were put within wrong genera (Attus, institutions and colleagues: Otago Museum, Dunedin (Brian , Salticus), many others are to be described. Patrick, Erena Barker, Simon Wylie); Auckland Museum In the past the efforts to provide a proper list of New Entomology Collection, Auckland (John Early); Canterbury Zealand Salticidae or to verify early descriptions were Museum, Christchurch; Lincoln University, Lincoln (Cor undertaken by Urquhart (1892), Parrott (1942) and Vink); Te Papa, Wellington (Phil Sirvid); Landcare Research Bryant (1935a, b) – all with limited success. NZAC (Trevor Crosby); Canterbury University, Christ- In 2000 we began a long-term project on church (Robert Jackson, Christchurch). The study was also and zoogeography of New Zealand Salticidae with the supplemented by our own field research. following goals: (1) to describe (redescribe) all the taxa available from the country, (2) to analyse the faunistic diversity for particular areas and habitats, (3) to RESULTS analyse the relationships between New Zealand and other faunas of the S-W Pacific, (4) to estimate the influ- Jumping – a model-group ence of Gondwanan heritage and post-Gondwanan iso- lation on New Zealand salticids and, in more distant Jumping spiders have long been considered a good future: (5) to reconstruct the zoogeographical history of model for zoogeographical research (Prószyñski 1976, Salticidae of Australia, and New Zealand. 1996; ¯abka 1990a, 1991b, 1993; Patoleta and ¯abka In this paper we provide general introductory com- 1999; ¯abka and Nentwig 2000). ments, our approach to the problem and remarks on salti- 1. The family is well defined, diverse (about 5000 species cid age, distribution, habitat preferences and dispersal. described) and its taxonomy is relatively well studied. 460 M. ŻABKA, S. D. POLLARD and M. ANSTEY

Figure 1. The beaches of New Zealand are littered with driftwood, showing the possibility of rafting. West coast, Harihari area. (Photo by M. ¯abka).

Figure 3. Dacrydium sp. is one of the Gondwanan relics. West coast, Harihari area. Figure 2. Tree fern forests are known to support (Photo by M. ¯abka). many endemic salticids. West coast, Harihari area. (Photo by M. ¯abka).

Figure 4. Mountain moss forests support New Zealand endemics. West coast, Harihari area. (Photo by M. ¯abka).

2. In geological scale the group seems old enough to New Zealand those of Australia and western Pacific analyse the impact of long-term processes. islands are the best known (e.g., Wanless 1988; Prószyñski 3. There is some data on dispersal of the family and 1992, 1996; ¯abka 1988a, 1990b, 1991a, 1994, 2000; Berry particular species. et al. 1996, 1997, 1998) making a good basis for this study. 4. The local/continental faunas are well distinguished. During the last 30 years a number of papers on differ- The age ent salticid genera and subfamilies have been published by Galiano, Logunov, Maddison, Prószyñski, Wanless, The salticid fossils are 30–50 milion years old Weso³owska, ¯abka, and others (for complete review see (Prószyñski and ¯abka 1980, 1983, Wanless 1984, Prószyñski 2002). Of all the faunas potentially influencing Wunderlich 1986, ¯abka 1988b) and the evidence for the ZOOGEOGRAPHY OF SALTICIDAE OF NEW ZEALAND 461

Figure 5. The river beds are important migration routes for spiders, e.g. for Bianor sp. West coast, Franz Joseph area. (Photo by M. ¯abka).

Figure 7. The mountain rainforests are amongst the most spectacular and beautiful on Earth. West coast, Harihari area. (Photo by M. ¯abka).

Figure 8. Manuka (Leptospermum scoparium) Figure 6. Some central and western mountain ranges were rapidly created during the last few forests resemble wet sclerophyll forests of million years, providing unique conditions for speciation. Mt Cook, 3764 m. (Photo by M. ¯abka). Australia, but, with few exception (e.g., single species of Holoplatys) they support different communities. Canterbury, Hinewai. M. Dispersal (Photo by M. ¯abka). To understand the distribution of the group, its dis- present-day genera is even younger (Cutler 1984, Wolff persal has to be considered. In our case rafting, bal- 1990). However, the diversity of the fossil taxa, their looning and human agency seem the most likely ways of affinities with modern salticids and the presence of dispersal. Some beaches of New Zealand, are covered Salticidae in New Jersey amber (Wayne with masses of driftwood. Although most of it is of local Maddison – pers. comm.) suggest that the family is origin, the specimens and egg-sacs of Australian much older and that the present-day genera might have Holoplatys, that can be found in crevices and under existed much earlier (Prószyñski and ¯abka 1983, bark, suggest long-distance rafting from Australia Cutler 1984). which is quite likely due to favourable sea currents. A 462 M. ŻABKA, S. D. POLLARD and M. ANSTEY mystery is the presence of “Marpissa marina” (related If we are going to search for southern faunistic relics, to Australian Lycidas) on pebbly beaches and rocky we have to consider the age of taxa, their taxonomic level shore cliffs. The strong retreats and (water-proof?) egg- and appropriate endemic biota. Any Gondwanan sacs that can be found under rocks and stones in the Salticidae to occur in New Zealand, must have existed at tidal zone, seem a perfect adaptation for rafting. least 60–80 million years ago. As the oldest known identifi- However, neither specimens, retreats nor egg-sacs of able genera are much younger, the search for Gondwanan this species have been observed on driftwood. If relics may seem pointless though the taxonomic diversity “Marpissa marina” does disperse by rafting, it seems of the oldest fossils suggests that the present-day genera to do it only within New Zealand as it has not been may have existed much earlier than we think today. recorded anywhere else. However, the question whether they have witnessed the Ballooning is another way of dispersal. Theoretically, break up of Gondwana remains with no answer. the size and favourable W→E winds nominate Australia The search for Gondwanan salticid relics has usual- the best source of ballooners. In contrast to other spider ly been a side-effect of studies on the taxonomy of trop- families (araneids, linyphiids, oxyopids), salticids con- ical and subtropical faunas. Such an approach could not stitute only a tiny fraction (about 1.8%) of araneoplank- have been successful because warm-climate-biota are of ton (Horner 1975; Salmon and Horner 1977, Greenstone more recent origin. The salticids of Australian et all. 1987; Blandenier and Fürst 1998) and as such are forest, for instance, show very high rather poor candidates for aerodispersal. It seems that (neo)endemism, but no evidence of Gondwanan heritage only Helpis minitabunda is an effective Australian bal- (¯abka 1990a, 2000). looner that reaches New Zealand and inhabits warmer So, where to look for Gondwanan relics? parts of North and South Islands. In this case human Theoretically, the old temperate biota such as agency may also contribute (Patoleta and ¯abka 1999). Nothofagus-, Podocarpus- or Dacrydium-forests, either in New Zealand or in , and The distance between Australia and New Zealand acts south Chile are to be considered. Our preliminary data as a fine sieve through which only few ballooners can from New Zealand and Australia (including Tasmania) pass. Also narrow niche specialists of Australia are (¯abka unpubl.) make us rather sceptical about southern poor candidates for successful colonisers of completely salticid relics. Even if they do exist in New Zealand, they different biota on the other side of Tasman Sea. will be difficult to identify as such because of lack of Of five candidate-species that may have arrived with appropriate southern faunas elsewhere to compare them humans, only Hasarius adansoni has been recorded with (some suggest southern Chile would be a good area on the North Island. to take a closer look). It is likely that in the majority of While rafting, ballooning or human agency are nec- temperate habitats, southern salticid taxa have been essary for newcomers to arrive, the spreading within replaced by modern neoendemics – as the example of the country can (further) be made by slow penetration. Australia seems to suggest (¯abka 1990a). River beds seem one of the possible routes. Species of A very important factor to be considered in under- Bianor, for instance, and representatives of some standing New Zealand salticid fauna is the young age endemic genera are found to migrate this way. (few million years) of central and western mountain “Marpissa” marina can spread along the sea shore. ranges (Richard Holdaway pers. comm.). As they were rapidly created, the various areas would have become isolated from each other quite recently (not much more NEW ZEALAND – A GONDWANAN ARC? than 1 million years BP) and this could explain close relationships between species within “Trite” auricoma New Zealand is well known for its flightless birds and T. minax [= planiceps]-groups. (extinct and living), tuatara, lack of native land mam- The geological history will, of course, have had an mals, diverse and archaic frog fauna, endemic inverte- impact on today’s salticid fauna. However, instead of a brates (including many endemic spider taxa – see desperate search for Gondwanan relics, we should Platnick 2001) and unique flora, and has been a labora- notice more obvious evidence of New Zealand “lonely tory-area for studies on island life, especially on the role voyage” of which high endemism is the best manifesta- of isolation, extinction and colonisation rates, specia- tion. Whether the endemics are also relics is a matter tion, endemism and human influence. for future study. Some 60–80 million years ago New Zealand drifted away from other parts of Gondwana and has remained in long-distance isolation ever since. The climatic and IS AUSTRALIA A FAUNISTIC SOURCE? habitat changes during this lonely voyage resulted in mass extinction, speciation and high endemism of fau- Australia is the nearest (though still distant), the nas and floras. largest and the most natural source of colonisers – quite ZOOGEOGRAPHY OF SALTICIDAE OF NEW ZEALAND 463 effective for other , including spiders 3. Two groups of genera predominate the taxonomic (Lehtinen 1980, 1996; Main 1981). As shown earlier composition, of which “Trite” auricoma and Trite minax (¯abka 1991b), the Australian taxa are highly endemic [=planiceps] are the best known representatives. because they evolved in unique biota. Consequently, 4. The relationships between New Zealand and they have become “prisoners” of their habitats (mostly Australian faunas are limited to some genera, which are Eucalyptus forests) of Australia, New Guinea and some highly diverse in Australia and have only single repre- adjacent islands. sentatives in New Zealand. The comparison of the faunas of temperate forests of 5. No Gondwanan salticid relics have been distin- New Zealand with those in , and guished in New Zealand. Post-Gondwanan evolution of Tasmania also shows limited similarities. Some genera, biota/taxa, especially those caused by modern orogenic highly diverse in Australian, e.g., Holoplatys or processes, seems to be responsible for close relation- Opisthoncus, are represented in New Zealand by single ships between genera within “Trite” auricoma and T. species and occur mostly in manuka (Leptospermum sco- minax [= planiceps]-groups. parium) and kanuka (Kunzea ericoides) forests which, to some extent, are the local equivalent of Eucalyptus forests of Australia (open character, similar bark struc- ACKNOWLEDGEMENTS ture). Other taxa, very diverse in Australia (Servaea, Adoxotoma, Arasia, Maratus and many others) have not The research was possible due to the Grant been found in New Zealand so far. Thus, the limited rela- 6PO4C01115 provided for M¯ by the Polish State tionships between New Zealand and Australian faunas is a Committee for Scientific Research. M¯ is grateful to the consequence of biota uniqueness, climatic limitations, and Director and staff of Canterbury Museum (Christchurch, space isolation of both lands (islands). New Zealand), and especially to the friends in Invertebrate Zoology who were most kind and co-opera- tive during his 3-month long visit there. Brian Patrick, OTHER RELATIONSHIPS Erena Barker, John Early, Phil Sirvid and Trevor Crosby – the curators of several New Zealand collections (listed Prószyñski (1996) analysed the salticids of the Pacific earlier) are acknowledged for providing material for Islands and found that only a few widely distributed gen- study. The project was also supported by Michael Gray era occurred also in New Zealand. Our studies support (Australian Museum, Sydney), Robert Raven his views. Single species of Trite miax [= planiceps]- group have expanded to some islands of Polynesia, ( Museum, Brisbane), Robert Jackson Melanesia and Micronesia (human agency?), at the same (Canterbury University, Christchurch) and Cor Vink time single wide-range genera (e.g., Bianor, Neon) have (Lincoln University, Lincoln). Jerzy Prószyñski (Museum their representatives in New Zealand. Rather mysteri- and Institute of Zoology, Polish Academy of Sciences, ous is the group of 10 (maybe more) species related to Warsaw) and Wanda Weso³owska (Wroc³aw University) Euophrys and Lycidas; their affinities are to be verified provided critical comments on the text. Our research (¯abka in prep.). The reasons for limited faunistic rela- would not have been possible without decades of arach- tionships between New Zealand and other Pacific archi- nological research carried out by the late Ray Forster pelagos seem the same as discussed for Australia. 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Received: February 8, 2002 Accepted: May 28, 2002