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Population Genetic Structures of Two New Zealand Stream Insects: Archichauliodes Diversus (Megaloptera) and Coloburiscus Humeralis (Ephemeroptera)

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Population Genetic Structures of Two New Zealand Stream Insects: Archichauliodes Diversus (Megaloptera) and Coloburiscus Humeralis (Ephemeroptera) NewHogg Zealand et al.—Population Journal of Marinegenetics and of New Freshwater Zealand Research, aquatic insects 2002, Vol. 36: 491–501 491 0028–8330/02/3603–0491 $7.00 © The Royal Society of New Zealand 2002 Population genetic structures of two New Zealand stream insects: Archichauliodes diversus (Megaloptera) and Coloburiscus humeralis (Ephemeroptera) IAN D. HOGG Keywords gene flow; allozymes; Ephemeroptera; PIA WILLMANN-HUERNER Megaloptera; biodiversity; dispersal MARK I. STEVENS Centre for Biodiversity and Ecology Research Department of Biological Sciences INTRODUCTION University of Waikato Identifying patterns of genetic variability within Private Bag 3105 species may be critical from a conservation Hamilton, New Zealand perspective as it is widely contended that email: [email protected] maintaining such diversity will facilitate both population and ecosystem-level responses to environmental change. However, concern is often Abstract We examined the population genetic targeted towards overall levels of variability, while structure (allozyme variation) of two New Zealand relatively little consideration is given to how the aquatic insects: Archichauliodes diversus (Walker) variability may be partitioned among populations (Megaloptera) and Coloburiscus humeralis (Walker) (i.e., genetic differentiation). For freshwater systems, (Ephemeroptera); and tested the hypothesis that spe- which often consist of naturally fragmented and cies with limited adult dispersal abilities would show isolated habitats, this may be particularly relevant high levels of genetic differentiation among (Hogg et al. 1998a). Specifically, interhabitat hydrologically isolated habitats. Both species were dispersal (and hence gene flow) may be restricted for characterised by low levels of within-population some species resulting in limited within-population genetic variability (e.g., Hexp = 0.06 and 0.04 for A. variability—an attribute potentially compounded by diversus and C. humeralis, respectively). However, the increasing anthropogenic fragmentation of genetic differentiation among sites was considerably aquatic habitats. greater for A. diversus relative to C. humeralis (FST The extent to which populations of a species be- = 0.57 and 0.03, respectively). A. diversus consisted come genetically differentiated will be determined of two genetically distinct groups in the North Island in part by: (1) the biological characteristics of the (I = 0.96), and a third group from the South Island species (e.g., dispersal abilities, geographic distribu- (I = 0.67). By contrast, allelic differences among C. tion); (2) selective pressures that have been imposed humeralis were minimal (I > 0.99, in all cases), and on local populations (Slatkin 1985); and (3) random appeared unrelated to geographic proximity or habi- genetic drift (Jackson & Resh 1992). However, tat type. We suggest that a combination of historic present-day patterns of population structure are also range changes and reproductive mechanisms, as well likely to have been influenced by the history of a as contemporary dispersal capabilities, may be re- particular species (e.g., genetic bottlenecks, breed- sponsible for the observed population genetic struc- ing systems). Accordingly, genetic assessments also tures of A. diversus and C. humeralis. Accordingly, need to consider the species’ past (e.g., Bossart & generalisations based on present-day dispersal abili- Prowell 1998; Driscoll 1998). Unfortunately, it is ties alone may be misleading particularly for island often difficult to separate the relative importance of habitats such as New Zealand where gene pools may present-day events relative to the past history of a already be restricted. species. New Zealand may provide an ideal opportunity to test hypotheses related to patterns of species and M01069; published 17 September 2002 genetic diversity. Specifically, it has a small land Received 26 July 2001; accepted 19 March 2002 mass (270 000 km2), wide latitudinal range (c. 12°), 492 New Zealand Journal of Marine and Freshwater Research, 2002, Vol. 36 1700 1750 1800 Fig. 1 Study area, showing loca- tions of sampling sites from 350 Northland (N1–4), Waikato (W1– N3 N2 4), central North Island (C1) and N1 South Island (S1), New Zealand. N4 North Island W4 W2,W3 W1 C1 400 South Island N 450 S1 300 km diverse topology, and well documented geologic somewhat restricted (e.g., Ephemeroptera, history (Suggate et al. 1978). Furthermore, New Trichoptera, Hemiptera). By contrast, studies of Zealand has been isolated in the south Pacific Ocean North American taxa have suggested con- since separation from Australia some 70 million siderably greater levels of differentiation even at years ago (Gibbs 1989). Accordingly, stream insects smaller spatial scales (e.g., Jackson & Resh 1992; with potentially limited dispersal abilities inhabiting Hughes et al. 1999). New Zealand’s highly fragmented landscape may To enhance our knowledge of the New Zealand serve as an ideal model for comparative studies fauna, we evaluated the population genetic structures assessing the relative contributions of present and of two widespread and endemic stream insects, past events. Archichauliodes diversus (Walker) (Megaloptera) Before beginning this study, little was known of and Coloburiscus humeralis (Walker) (Epheme- New Zealand’s stream insects, although a single roptera). By selecting species with limited dispersal study has since been published (Smith & Collier capabilities from discrete habitats (i.e., different 2001). Comparatively more is known of the streams and catchments), we reasoned that present Australian fauna (e.g., Schmidt et al. 1995; Bunn & day gene flow should be minimal, and that Hughes 1997; Hughes et al. 1998, 1999, 2000). populations should show significant genetic These studies have generally suggested limited differentiation among locations. As we show, results levels of genetic differentiation among populations, for C. humeralis are contrary to these expectations although taxonomic coverage still remains and require an alternative explanation. Hogg et al.—Population genetics of New Zealand aquatic insects 493 METHODS Collection of samples Study species Mature (late instar) larvae of A. diversus and C. humeralis were collected between February and Archichauliodes diversus , the only New Zealand October 1998. Larvae were caught with dipnets (250 megalopteran, has a predacious and almost entirely µm) by kick sampling and by turning and washing aquatic (larval) existence (Devonport & Winter- large stones to dislodge individual animals. The bourn 1976; Harding & Winterbourn 1993). contents of the nets were then placed into sorting Prolonged larval development (c. 18 months–3 trays where the study organisms were removed and years), long and asynchronous emergence periods counted. Individuals were placed in plastic sampling (Hamilton 1940; Devonport & Winterbourn 1976), jars containing stream water and transported back to and obligate diapause of late-laid eggs (Edwards the laboratory on ice. When fieldwork extended over 1986) have been suggested. Adults are non-feeding several days, individual animals were frozen and live from 6–10 days (Hamilton 1940). Their immediately in liquid nitrogen in the field. In the flight is slow and clumsy (Edwards 1986) and laboratory, the samples were stored at –75°C in individuals manage only short distances over streams individual 1.5 ml microcentrifuge tubes until needed at very low height because of their heavy bodies and for electrophoretic analysis. large, awkward wings (5–8 cm wing span). The mayfly, Coloburiscus humeralis (Oligo- Genetic analysis neuriidae) exhibits a filter-feeding, aquatic larval stage (Tan 1961), with life history ranging from All study organisms were morphologically con- univoltine in the warmer, northern warmer regions, firmed using Winterbourn & Gregson (1989). To to 18–27 months in lake outlet streams and cool, evaluate genetic differences within and among forested streams (Tan 1961; Norrie 1969). Although populations, we used cellulose acetate electrophore- no specific information is available for adult C. sis following techniques described by Hebert & humeralis, mayflies are known to be short-lived as Beaton (1993) and Larose & Hogg (1998). adults (e.g., 10–30 min in the burrowing mayfly Initially, enzyme systems showed very poor Dolania americana) (Sweeney & Funk 1991). resolution for A. diversus with smudged bands However, some ovoviviparous species have been complicating reliable identification of allelic reported to live up to 14 days (Brittain 1982). variation. Since it was suspected that the gut contents of this predatory organism could be confounding Study sites band resolution, trial runs were conducted using Ten sampling sites were selected to examine the different body parts (e.g., head, thorax, abdomen). genetic structure of A. diversus and C. humeralis Although banding patterns were the same, best (Fig. 1). Four sites within the Waikato region (W1– results were obtained when only the heads were W4) were in relatively close proximity to each other, used. Accordingly, individual heads were separated by 1.5–18 km. The Northland sites (N1– homogenised in 20–30 µl of a grinding buffer N4) were separated by larger distances (c. 50– containing distilled water (100 ml), NADP (10 mg), 75 km) and were c. 200–300 km away from the b-mercaptoethanol (100 µl), and detergent Tween 80 Waikato sites. The maximum distance occurred (100 µl) (Richardson et al. 1986). Samples were then between the uppermost
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