Allozyme characterization of Hogna species (Araneae, Lycosidae) of the Galapagos Archipelago.
The Galapagos Archipelago consists of 13 large islands and a great number of islets and rocks, all of volcanic origin (Fig. 1). The southeastern islands are the oldest (3-5 million years) while the northern and western islands are the youngest (< 0.7 million years) (Simkin 1984).
Due to geographic isolation, many endemic animal (e.g., Darwin's finches, giant tortoises, lava lizards, mockingbirds) and plant (e.g., Opuntia cacti, Scalesia trees) groups have radiated. Evolutionary research on these islands has mainly focused on vertebrate species such as Darwin's finches, giant tortoises, lava lizards, mockingbirds, and on plant species such as Opuntia cacti and Scalesia trees (Grant 1981; Fritts 1984; Snell et al. 1984; Stern & Grant 1996; Rassmann et al. 1997; Cacone et al. 2002). Speciation patterns of invertebrates have been, in contrast, much less studied, and only recently have genetic studies been conducted on Coleoptera such as the tenebrionid Stomium (Finston & Peck 1995), the chrysomelid Nesaecrepidia (Verdyck & Desender 1999), the carabid Calosoma (Desender & Verdyck 2000; Verdyck et al. 2003, 2004), the weevil Galapaganus (Sequeira et al. 2000, 2008), and the land snail genus Bulimulus (Parent & Crespi, 2006). Genetic studies on Galapagos spiders are presently lacking, while such studies in other locations have revealed adaptive radiations on other archipelagos (e.g., Hawaian Tetragnatha spiders, Gillespie 2004; Hawaiian Dysdera spiders, Arnedo 2001).
Previous studies on the spider genus Hogna, the only wolf spider genus occurring on the archipelago (Maelfait & Baert 1986; Baert & Maelfait 1997), revealed that this genus consists of several closely related, or even cryptic, species. Based on somatic and small genital differences, a total of seven morpho-species is suggested with a distinct distribution on the islands (Baert et al. unpublished data) (Fig. 1). At least three groups of morpho-species can be distinguished that differ distinctly.
A first group of ecologically and morphologically similar morpho-species occur at higher altitudes on the islands in the pampa vegetation zone and are hereafter referred to as "high elevation species." Based on differences in morphology of the genital organs, different species can be distinguished: Hogna species 1 (H1), living on both southern volcanoes Cerro Azul and Sierra Negra of Isabela, one of the youngest islands, Hogna species 4 (H4) which occurs on islands of intermediate age (Santa Cruz, Santiago, and Volcan Alcedo of Isabela), and Hogna species 2 (H2) which occurs on the oldest island of San Cristobal.
A second group, hereafter referred to as "coastal dry species," lives in the dry arid zone along the coast in vegetated dunes and in the Opuntia cactus zone. These morpho-species can only be found on the oldest islands of San Cristobal (Hogna species 5 (H5)), Espariola (Hogna species 7 (H7)), and Santa Cruz (Hogna species 6 (H6)). The San Cristobal species (H5) is mostly found in the depressions overgrown with sea grass (Sporobolus virginicus) located behind the shore or on low vegetated dunes, while H7 is found in tall vegetation of the dry arid zone directly adjacent to the littoral zone, but never in the adjacent depressions with salt grass. The Santa Cruz species (H6) is found in the Opuntia cactus zone in between dune and pure rocky soil.
[FIGURE 1 OMITTED]
The third group comprises populations of the generalist species Hogna species 3 that lives in saline habitats along the coast (salt marshes, bays), along permanent pools (e.g., El Chato on Isla Santa Cruz) and in permanent wetlands below 600 m of altitude (e.g., Los Gemelos on Isla Santa Cruz). Scattered populations can also be found above the vegetation inversion zone in wet conditions during El Nino years (characterized by very heavy rainfall giving rise to temporary pools) (Baert & Maelfait 2000). They reach, however, their highest densities in the salt marshes. All these populations have very similar genital organs and are at present interpreted as belonging to a single species. It is widespread over the whole archipelago, with the exception of the northern island Pinta and the southeastern island Espannola (Baert & Maelfait 1997).
In this paper, we test whether the separation of this genus into seven morpho-species on the Galapagos is justified. By means of cellulose acetate gel electrophoresis, wherein 8 allozyme loci (FUM, G6PDH, GOT, IDH, LDH, MPI, PGI, and PGM) were studied, we investigate whether the genetic variation among species is larger compared to the variation among populations within species and indicative of reproductive isolation among the species.
Sampling collection.--In the period between 1996 and 2002, we sampled a total of 43 known Hogna populations (see Table 1) from 9 islands (Santa Cruz, Isabela Vokan Sierra Negra, Isabela Vokan Cerro Azul, San Cristobal, Floreana, Rabida, Genovesa, Bartolome, Santiago, and Espannola) and seven morpho-species. In three localities, the high elevation species occurred sympatrically with H3, [e.g., Cerro Gavillan (populations 40 & 41), El Junco (populations 1 & 2), and Los Gemelos (population 23) (Fig. 1)]. Populations of H4 occurring on the tops of Isabela and Santiago were preserved in ethanol and could therefore not be included in this allozyme study.
Individuals were caught by hand, mostly at sunset with an electric torch worn on the forehead. They were stored and transported in a Taylor-Wharton cryogenic shipper saturated with liquid nitrogen. In the laboratory, the material was stored in an ultra-cold freezer at -80[degrees] C. The aim was to investigate at least 40 individuals for each population if possible. In some localities, their densities were so low that this number could not be reached. Some localities were sampled several times but in different years. Voucher specimens are deposited at the Royal Belgian Institute of Natural Sciences.
Allozymes.--Parts of the legs were homogenized in distilled water for performing the cellulose acetate gel electrophoresis, following the procedures of Hebert & Beaton (1989). Eight enzymes (9 loci) were tested for polymorphism: fumarate hydratase (FUM), aspartate minotransferase (GOT1, GOT2), isocitrate dehydrogenase (IDH),lactate dehydrogenase (LDH), mannose phosphate isomerase (MPI), phosphoglucose isomerase (PGI), phosphoglucomutase (PGM), and 6-phosphogluconate dehydrogenase (6PGDH).
Allele frequencies were obtained for each population and species. Deviations from Hardy-Weinberg equilibrium were tested by means of an exact test. Genetic divergence between populations and species were estimated based on Nei's unbiased genetic distance (Nei 1978). Based on this distance metric, divergence among populations within species was compared with the distance among populations of different species. These analyses were performed with the computer packages TFPGA (Miller 1997) and GenAlEx (Peakall and Smouse 2006). Genetic distances were visualized by means of principal component analysis (PCA), designed for ordination of allelic frequency data, by means of the computer package PCA-Gen (Goudet 1999).
Allelic variation and heterozygosity were very low within each species, but differed clearly among species, with one or a few alleles that were fixed within the morpho-species. The low genetic variability among populations within species, compared to the variability among species, is clearly depicted when genetic distances are compared among populations (Table 2). The genetic distance between populations belonging to the same morpho-species ranged from 0 to maximum 0.031 (H3), demonstrating that the allele frequencies of the different populations within a given morpho-species were highly similar. Differences in allele frequency of populations belonging to a different morpho-species were, in contrast, considerably higher and ranged from 0.118 to 2.277. The smallest genetic distances were observed between H2 ("high elevation" San Cristobal) and H7 ("coastal dry" Espanola) and between H5 ("coastal dry" San Cristobal) and H7.
Allele frequencies for all loci were near to fixation for almost all morph-species (Table 3). None of the species comparisons resulted in fixation of the same alleles, and each morpho-species was, therefore, characterized by a unique allele combination.
The low differences among conspecific populations compared to differences among morpho-species were also obtained from PCA ordination of the different populations (Fig. 2). The first three axes explained 74.24%, 15.06%, and 4.06% of the total allelic variation respectively. Along the first axis, the H3 populations are clearly separated from the other morpho-species. The position of the remaining species along this axis and the second axis corresponded to their geographic position and age of the islands rather than to their habitat preference. The three species of the oldest islands--H7 from Espafiola and H2 and H5 from San Cristobal--were clustered in the PCA, followed by H4 and H6 from Santa Cruz and H1 from the volcanoes Cerro Azul and Sierra Negra on Isabela.
[FIGURE 2 OMITTED]
Our results show that there is a clear and very high degree of genetic divergence between the previously defined morpho-species. Moreover, genetic divergence among populations within species was much lower (Table 2, Fig. 2). These two findings indicate that these morpho-species likely represent distinct reproductively isolated species.
Although the validity of allozymes for phylogenetic inferences is questionable since the historical and genealogical relationship between the different alleles remains unknown (Lowe et al. 2004), a few suggestions concerning historical patterns of divergence can be deduced.
First, these results suggest that Hogna speciation on the Galapagos is likely due the combined effect of geographic isolation and ecological specialization within the different climatological and vegetation zones present on the different islands. Except for H3, species from the same or proximate island tend to be genetically more similar to each other. Species living on the same islands but in different habitats are, however, genetically fixed for a few loci, clearly indicating a lack of gene flow and hence strong reproductive isolation. Combining these results suggests that ecological specialization on the islands Santa Cruz and San Cristobal occurred repeatedly in association with speciation events rather than a diversification of a habitat adapted lineage with secondary colonization of the specialized forms to the different islands. According to the second scenario, species living in the same habitat would then be expected to be genetically more similar to each other.
Similar patterns of species diversification in terms of geographic position and ecological specialization have been confirmed by more thorough genetic analyses of Tetragnatha spiders from Hawai (Gillespie 2004). In the Galapagos, this speciation pattern has been observed in other terrestrial invertebrates such as weevils and snails (Parent & Crespi 2006; Sequiera et al. 2008).
Whether the generalist species H3 can be regarded as closely related to the ancestral species, as suggested by Baert & Maelfait (2000), however, cannot be confirmed by these data. The smaller genetic distance between the generalist H3 and the species from younger islands (H1) compared to those of older islands (H2, H5, and H7) is in accordance with this hypothesis. Surprisingly however, H3 showed a very low degree of genetic variation within populations. Moreover, distant populations as well as populations living in different habitats all appeared to be genetically very homogenous. These observations suggest that this generalist and apparently highly dispersive species may have colonized the archipelago independently.
The results can only be interpreted as preliminary as they are based on allozyme data and only a few loci were scored. Moreover, the selective neutrality of allozymes has often been questioned. Our ongoing work aims to add more variable loci such as mitochondrial DNA (Cytochrome Oxidase I) so that more well founded phylogenetic inferences can be made. Also, future work should include Hogna species from the South American mainland to better understand the phylogenetic relationships between the species and the colonization history of Hogna in Galapagos.
Excellent cooperation and field logistic support were provided by the Charles Darwin Research Station (CDRS, Isla Santa Cruz, Galapagos, Ecuador), the directors F. Koestner, G. Recek, D. Evans, C. Blanton, R. Bensted-Smith, M. Cifuentes and their staff; the Galapagos National Park Service (SPNG Superintendents M. Cifuentes, IR. H. Ochoa, F. Cepeda, A. Izurieta, and E. Cruz), Department of Forestry, Ministry of Agriculture of Ecuador; TAME airline kindly issued a reduced price for travel tickets. Our investigations and field work were financially supported by (1) BELSPO (former Belgian DWTC), (2) the Fund for Scientific Research (FWO Vlaanderen; research project G.0202.06), and (3) the Leopold III Foundation. Help in the field was provided by K. Desender, L. Roque, and P. Verdyck. Help with electrophoresis was provided by K. Desender, K. Loosveldt, and V. Versteirt. Constructive comments on a previous version were given by Marshal Hedin and an anonymous referee.
Manuscript received 27 November 2007, revised 15 July 2008.
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Leon Baert: Royal Belgian Institute of Natural Sciences, Entomology Department, Vautierstraat 29, 1000 Brussels, Belgium. E-mail: firstname.lastname@example.org
Frederik Hendrickx: Royal Belgian Institute of Natural Sciences, Entomology Department, Vautierstraat 29, 1000 Brussels, Belgium; Terrestrial Ecology Unit, Biology Department, Ghent University, K.L. Ledeganckstraat 35, 9000 Gent, Belgium
Jean-Pierre Maelfait: Research Institute for Nature and Forest (INBO), Kliniekstraat 25, 1070 Brussels; Terrestrial Ecology Unit, Biology Department, Ghent University, K.L. Ledegnackstraat 35, 9000 Gent, Belgium
Table 1.--List of the 43 sampled Hogna populations on Galapagos during the years 1996, 1998, 2000, and 2002. Situation and number of caught specimens per sample. Abbreviations: SCB = Isla San Cristobal; ESP = Espanola; GEN = Genovesa; SCZ = Santa Cruz; FLO = Floreana; BAR = Bartolome; RAB = Rabida; SAN = Santiago; ISN = Isabela, Volcan Sierra Negra; ICA = Isabela, Volcan Cerro Azul. Code Island Locality Vegetation zone 1 SCB El junco pampa 2 SCB El junco pampa 3 SCB Cerro San Joaquin pampa 4 SCB Punto Baso dune 5 SCB Punto Baso (fregat nesting) dune 6 SCB Punto Baso (Sesuvium) littoral zone 7 SCB Caleta de la Tortuga littoral zone 8 SCB La Loberia littoral zone 9 SCB Caleta Sapho (Spirobolus) salt grass 10 ESP Punta Cevallos dry arid zone 11 ESP Bahia Gardner dry arid zone 12 ESP Isla Gardner dry arid zone 13 GEN Lago Arcturus littoral zone (lagoon) 14 SCZ Laguna Andreas littoral zone (lagoon) 15 SCZ Bowdich littoral zone (lagoon) 16 SCZ El Garapatero littoral zone (lagoon) 17 SCZ Las Palmas dry arid zone 18 SCZ Meteo Station littoral zone (lagoon) 19 SCZ Bahia Tortuga littoral zone (lagoon) 20 SCZ Playa Bachas littoral zone (lagoon) 21 SCZ El Chato around permanent pool 22 SCZ El Carmen temporary pool (El Nino) 23 SCZ Los Gemelos, open pampa 24 SCZ Los Gemelos, Scalesia Scalesia forest 25 SCZ Media Luna pampa 26 SCZ Tss ML & Cpunt pampa 27 SCZ Cerro Crocker pampa 28 FLO Punta Cormoran littoral zone (lagoon) 29 FLO Finca Cruz pampa 30 FLO Highland pampa 31 BAR littoral zone (lagoon) 32 RAB littoral zone (lagoon) 33 SAN Playa Espumila littoral zone (lagoon) 34 SAN Aguacate transition zone (El Nino) 35 SAN La Central pampa (El Nino) 36 SAN Jaboncillo pampa (El Nino) 37 ISN Laguna de Villamil littoral zone (lagoon) 38 ISN Top pampa 39 ICA Caleta Iguana littoral zone/dry arid zone 40 ICA Cerro Gavilan pampa 41 ICA Cerro Gavilan pampa 42 ICA 1100m dry arid zone (El Nino) 43 ICA Top dry arid zone (El Nino) Annual total no. specimens Morpho- Locality Elevation species 1996 El junco 675m H3 8 El junco 675m H2 3 Cerro San Joaquin 700m H2 Punto Baso 5m H5 Punto Baso (fregat nesting) 5m H5 Punto Baso (Sesuvium) 1m H3 Caleta de la Tortuga 5m H3 La Loberia 1m H3 Caleta Sapho (Spirobolus) 1m H5 28 Punta Cevallos 2m H7 Bahia Gardner 1m H7 Isla Gardner 2m H7 Lago Arcturus 60m H3 Laguna Andreas 1m H3 35 Bowdich H3 El Garapatero H3 Las Palmas H6 Meteo Station 5m H3 Bahia Tortuga 1m H3 Playa Bachas 1m H3 El Chato H3 El Carmen 350m H3 Los Gemelos, open 570m H3 Los Gemelos, Scalesia H3 Media Luna 600m H3 Tss ML & Cpunt H4 Cerro Crocker 875m H4 Punta Cormoran 1m H3 Finca Cruz 200m H3 Highland 350m H3 1m H3 1m H3 Playa Espumila 1m H3 Aguacate 500m H3 La Central 700m H3 Jaboncillo 820m H3 Laguna de Villamil 1m H3 41 Top H1 42 Caleta Iguana 2m H3 Cerro Gavilan 700m H3 Cerro Gavilan 700m H1 1100m 1100m H3 Top 1530m H3 Annual total no. specimens 157 Locality 1998 2000 2002 El junco 36 El junco 44 Cerro San Joaquin 46 Punto Baso 41 Punto Baso (fregat nesting) 36 Punto Baso (Sesuvium) 27 Caleta de la Tortuga 10 La Loberia 42 Caleta Sapho (Spirobolus) 55 Punta Cevallos 40 Bahia Gardner 31 Isla Gardner 41 Lago Arcturus 18 Laguna Andreas 42 40 Bowdich 31 El Garapatero 58 22 Las Palmas 28 Meteo Station 40 Bahia Tortuga 41 27 53 Playa Bachas 43 40 44 El Chato 40 40 El Carmen 42 Los Gemelos, open 43 Los Gemelos, Scalesia 47 40 Media Luna 51 37 27 Tss ML & Cpunt 13 Cerro Crocker 42 23 47 Punta Cormoran 40 Finca Cruz 15 Highland 50 31 38 Playa Espumila 45 40 Aguacate 40 41 La Central 38 40 Jaboncillo 40 40 Laguna de Villamil Top Caleta Iguana 32 Cerro Gavilan 12 Cerro Gavilan 30 1100m 11 Top 30 Annual total no. specimens 606 549 926 Table 2.--Genetic distance data for seven Hogna morpho-species on the Galapagos. Above diagonal: genetic distances between the different morpho-species when allele frequencies of the different populations of each morpho-species were pooled; diagonal: minimum and maximum genetic distance between populations of the same morpho-species; below diagonal: minimum and maximum genetic distance between populations of different morpho-species. H1 H2 H3 H4 H1 0.012 1.711 1.026 0.449 H2 1.647-1.811 0.0016 2.059 0.587 H3 0.910-1.040 1.614-2.154 0.000-0.031 1.481 H4 0.435-0.483 0.577-0.600 1.230-1.504 0.0001 H5 1.179-1.281 0.260-0.275 1.213-1.503 0.575-0.584 H6 1.084-1.097 1.123-1.182 1.083-1.235 0.602-0.607 H7 1.645-1.841 0.122-0.146 1.727-2.277 0.798-0.811 H5 H6 H7 H1 1.218 1.089 1.719 H2 0.267 1.151 0.131 H3 1.474 1.090 2.193 H4 0.577 0.603 0.800 H5 0-0.0001 0.817 0.125 H6 0.813-0.822 -- 1.119 H7 0.118-0.138 1.121-0.122 0.000-0.013 Table 3.--Allele frequency data for seven Hogna morpho-species on the Galapagos. H1 H2 H3 H4 H5 H6 Npop 2 2 30 2 3 1 Nind 72 91 1632 144 112 28 FUM 1 0.0000 0.0000 0.0000 0.0035 0.0000 1.000 2 0.9931 0.0000 0.0000 0.9896 0.0000 0.0000 3 0.0000 0.9890 0.0000 0.0035 1.0000 0.0000 4 0.0069 0.0110 1.0000 0.0035 0.0000 0.0000 GOT1 1 0.0000 0.0165 0.0000 0.0000 0.0000 0.0000 2 0.0278 0.9560 0.0018 1.0000 1.0000 1.0000 3 0.9722 0.0220 0.9982 0.0000 0.0000 0.0000 4 0.0000 0.0055 0.0000 0.0000 0.0000 0.0000 LDH 1 1.0000 0.0000 0.9997 0.0035 0.0000 0.0000 2 0.0000 0.0000 0.0000 0.0069 1.0000 0.0000 3 0.0000 1.0000 0.0003 0.9861 0.0000 0.0000 4 0.0000 0.0000 0.0000 0.0035 0.0000 1.0000 G6P3H 1 0.0000 1.0000 0.0003 0.0000 0.0000 0.0000 2 1.0000 0.0000 0.9994 0.9931 1.0000 1.0000 3 0.0000 0.0000 0.0003 0.0069 0.0000 0.0000 MPI 1 0.0000 0.0714 0.9997 0.0035 0.0045 0.0000 2 0.0000 0.9286 0.0003 0.0070 0.9955 0.0000 3 1.0000 0.0000 0.0000 0.9860 0.0000 0.0000 4 0.0000 0.0000 0.0000 0.0035 0.0000 1.0000 PGI 1 0.0000 0.0330 0.0003 0.0139 0.0000 0.0179 2 0.0069 0.8791 0.9988 0.9549 0.9911 0.9286 3 0.0139 0.0824 0.0009 0.0174 0.0089 0.0536 4 0.9583 0.0000 0.0000 0.0139 0.0000 0.0000 5 0.0208 0.0055 0.0000 0.0000 0.0000 0.0000 PGM 1 0.0000 0.0275 0.9969 0.0035 0.0000 0.0536 2 0.0278 0.9341 0.0031 0.0000 0.9688 0.0000 3 0.9722 0.0330 0.0000 0.9931 0.0313 0.9464 4 0.0000 0.0055 0.0000 0.0035 0.0000 0.0000 GOT2 1 0.0625 0.0495 0.9991 0.0069 0.0089 0.0000 2 0.9375 0.9286 0.0009 0.9861 0.9911 0.9286 3 0.0000 0.0220 0.0000 0.0069 0.0000 0.0714 IDH1 1 0.0000 0.0000 0.9694 0.0104 0.0000 1.0000 2 0.5000 1.0000 0.0306 0.9896 1.0000 0.0000 3 0.5000 0.0000 0.0000 0.0000 0.0000 0.0000 H7 Npop 3 Nind 153 FUM 1 0.0000 2 0.0000 3 1.0000 4 0.0000 GOT1 1 0.0033 2 0.9967 3 0.0000 4 0.0000 LDH 1 0.0000 2 1.0000 3 0.0000 4 0.0000 G6P3H 1 1.0000 2 0.0000 3 0.0000 MPI 1 0.0000 2 1.0000 3 0.0000 4 0.0000 PGI 1 0.0000 2 0.9412 3 0.0588 4 0.0000 5 0.0000 PGM 1 0.0000 2 0.8170 3 0.0065 4 0.1765 GOT2 1 0.0000 2 1.0000 3 0.0000 IDH1 1 0.0000 2 1.0000 3 0.0000
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|Author:||Baert, Leon; Hendrickx, Frederik; Maelfait, Jean-Pierre|
|Publication:||The Journal of Arachnology|
|Date:||May 1, 2008|
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