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Journal of Natural History Vol. 42, Nos. 5–8, February 2008, 491–511
Biogeographic patterns of tenebrionid beetles (Coleoptera, Tenebrionidae) on four island groups in the south Aegean Sea Apostolos Trichasa*, Anastasia Lagkisb, Kostas A. Triantisa,b, Nikos Poulakakisa and Maria Chatzakia a Natural History Museum of Crete, University of Crete, Crete, Greece; bBiology Department, University of Crete, Crete, Greece
New records on Tenebrionidae from four island groups of the southern Aegean (Santorini, Astypalaia, Nisyros and Kalymnos, with their satellite islets – 26 islands in total) allowed investigation of the biogeography of the taxon in the area. All tenebrionid records from small islets are new, with the exception of those from Pserimos and the satellite islets of Santorini. Chorological and nestedness analyses, species-area relationships and various measures of faunal similarity based on binary and quantitative data matrices were applied in order to explore biogeographic patterns. The Anatolian and east Mediterranean faunal elements dominate on the island groups of Kalymnos and Nisyros, with a considerable decrease on Astypalaia and Santorini, where circum-Mediterranean elements prevail. Analyses of faunal similarities clustered together the well defined island groups of Santorini and Astypalaia, while Kalymnos and Nisyros groups were clustered together rather vaguely. Astypalaia and Santorini were also the most nested island groups in the studied area. The biogeographic position of Astypalaia within the central Aegean islands is well supported by the data. Keywords: Aegean; Tenebrionidae; small islands; species-area; nestedness; biogeography; Santorini; Astypalaia; Kalymnos; Nisyros
Introduction The Aegean Sea is a marginal basin between Greek and Turkish peninsulas, penetrating the European continent from south-east and bounded toward the eastern Mediterranean Sea by the Hellenic Island Arc. The latter represents a small but geodynamically extremely active island arc, part of the Alpine-Himalayan orogenic system (Le Pichon and Angelier 1979; Angelier et al. 1982). The archipelago within the Aegean area consists of 7850 islands (large and small islands, islets and bare rocks above the sea-surface). Of these, 410 islands exceed 1 km2 in area. With the exception of the islands of the volcanic arc (Aegina, Poros, Milos, Santorini, Kos, Gyali and Nisyros), the majority of the central Aegean islands share a common geologic and palaeogeographic history, lying on the well defined Aegean plate, being in fact the mountain tops of the ancient land called ¨ ga¨is (Philippson 1898; Angelier et al. 1982; Kissel and Laj 1988; Ten Veen and A Meijer 1998). The Dodecanese islands represent a chain of islands mainly continental in origin. Starting in the late 1970s and increasing rapidly in the late 1990s and subsequently, biogeographic research in the Aegean area has provided good information on several animal groups such as land snails, isopods, centipedes, *Corresponding author. Email:
[email protected] ISSN 0022-2933 print/ISSN 1464-5262 online # 2008 Taylor & Francis DOI: 10.1080/00222930701835472 http://www.informaworld.com
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492 A. Trichas et al. tenebrionid beetles, butterflies, reptiles and amphibians (Mylonas 1982; Sfenthourakis 1996a; Trichas 1996; Foufopoulos and Ives 1999; Welter-Schultes and Williams 1999; Cameron et al. 2000; Dennis et al. 2000; Fattorini 2002; Chatzaki 2003; Chatzimanolis et al. 2003; Simaiakis et al 2005; Triantis 2006, and references therein). The huge number of islands with a broad range of size, the accessibility at least of the inhabited ones, the transitional biogeographic character of the area (combining Anatolian-East Mediterranean and Balkan-European elements) and the progress of knowledge on several taxa, are some of the causes for the increasing scientific interest in the Aegean archipelago. Despite the increasing number of studies, research effort on very small and less accessible groups of islands or islets are still embryonic, restricted to just a few studies in the Aegean area (for examples see Botsaris 1996; Panitsa and Tzanoudakis 2001; Bergmeier and Dimopoulos 2003; Triantis et al. 2005a; Panitsa et al. 2006). The principal aim of the present research is to contribute new data on the tenebrionid fauna of certain small and less known Aegean islands and islets, and to investigate the biogeographic relationships of these small island groups. Three out of the four island groups analysed in the present study (Astypalaia, Nisyros and Kalymnos) were lacking satisfactory information regarding their tenebrionid fauna (see Fattorini 2002; Hausdorf and Henning 2005). Materials and methods Study area Four small archipelagos in the south Aegean area were studied: (1)
(2)
(3)
Santorini island group, the southernmost Cycladic island aggregate. The present-day crescent shape of the islands is a consequence of the activity of the volcano in the past two million years. Except for a small part of original land, all the islands belonging to the Santorini group were subsequently formed by volcanic eruptions (Druitt et al. 1999). Sampling took place on the main volcanic island of Thira or Santorini (75.74 km2) and on Thirasia (9.23 km2). There are three more volcanic islets: Palia Kameni (0.52 km2), Aspronissi or Aspro (0.13 km2) and Nea Kameni (the most recent volcanic islet, not considered in the present study) (Figure 1). Astypalaia island group, a small and isolated complex, with Astypalaia being the largest out of 21 small islands. It is the only island formation between the Cycladic archipelago and the Dodecanese. The age and the degree of isolation of Astypalaia are obscured, with parts of the archipelago being not older than 15,000 to 22,000 years (Lambeck 1996; Perissoratis and Conispoliatis 2003) and other parts dating back to the upper Miocene (Dermitzakis and Papanikolaou 1981; Dermitzakis 1990). Sampling took place on the main island of Astypalaia (95.87 km2) and on eight islets: Kounoupi (1.44 km2), Agia Kyriaki (0.25 km2), Koutsomytis (0.45 km2), Ofidousa (1.91 km2), Pontikousa (0.97 km2), Mikro Fokionisi (0.09 km2), Megalo Fokionisi (0.57 km2) and Fteno (0.02 km2) (Figure 2). Kalymnos island group, which belongs to the Dodecanese and lies between Leros and Kos, very close to the Turkish coast. Most of the Kalymnos islands were disconnected very recently (,20,000 years) from the nearby coast, the whole island group being part of a landmass that until 10,000
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Journal of Natural History
Figure 1. Map of the island group of Santorini, its relative position in the Aegean and the sampling stations (crosses5pitfall trapping stations).
(3)
(4)
years ago also included the islands of Leros and Arkoi (Van Andel and Shackleton 1982; Lambeck 1996; Perissoratis and Conispoliatis 2003). It seems that Telendos and Kalavros are the most recent islets, being separated in historical times (,2000 years, Papageorgiou 2001). Sampling took place on the main island of Kalymnos (110.8 km2) and on seven islets: Agia Kyriaki (0.15 km2), Pserimos (14.63 km2), Plati (0.72 km2), Kalavros (0.28 km2), Nera (0.5 km2), Telendos (4.65 km2) and Sari (0.03 km2) (Figure 3). Nisyros island group, one of the newest volcanoes in the volcanic arc, not older than 200,000 years according to Vougiokalakis (1993). The caldera has taken its modern shape even more recently (30,000–15,000 years ago, Papanikolaou et al. 1991; Hardimann 1999). Sampling took place on the main island of Nisyros (41.47 km2) and on four islets: Gyali (4.54 km2), Pergousa (1.16 km2), Kandeliousa (1.36 km2) and Pacheia (1.16 km2) (Figure 4).
The area of each island (measured in km2) was provided by the 1:50,000 digitized maps of the Greek Army Geographic Service, using ESRI ArcGIS Desktop (Redlands, California, USA). Sampling Sampling was carried out by pitfall trapping and hand collections (see Figures 1–4 for the sampling stations). Unbaited traps (plastic caps of 12 cm height and 9.5 cm
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494 A. Trichas et al.
Figure 2. Map of the island group of Astypalaia, its relative position in the Aegean and the sampling stations (dots5hand collections, crosses5pitfall trapping stations).
opening, 15 caps per station containing ethylene glycol as preservative agent) were set in the period between 24 April and 3 May 2005 and collected in the period between 5 and 13 June 2005. Sampling sites represented the main habitat types of each island, therefore four to six sampling stations were established on each of the main islands, and one to three on the islets. Data analyses Tenebrionidae were identified to species (and subspecies for a few, widely accepted Aegean taxa, see Fattorini et al. (1999) for further comments) and all the presence/ absence data were recorded in binary matrices together with data from the literature. Literature records and species ranges were taken from Gridelli (1929), Koch (1935, 1948), Ku¨hnelt (1965), Ardoin (1976), Dajoz (1976), Grimm (1981), Fattorini et al. (1999) and Lo Cascio and Scupola (2003) (see Appendix). All collected material has been deposited in the Natural History Museum of Crete. In the chorological analysis the distribution ‘‘types’’ given by Fattorini et al. (1999) were followed (see also Leo and Fattorini 2002; Fattorini and Fowles 2005). All taxa were classified into the following biogeographic groups: (ANA): Anatolian taxa (distributed in the Anatolian peninsula and for some also at the Levantine, but not present at the northern coasts of Africa or the Balkans); (BAL): Balkan taxa (with distributions up to the northern parts of the Balkan peninsula, sometimes reaching the northern coasts of Anatolia and the Aegean islands to the south);
495
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Figure 3. Map of the island group of Kalymnos, its relative position in the Aegean and the sampling stations (dots5hand collections, crosses5pitfall trapping stations).
(WID): circum-Mediterranean, east Mediterranean, south European and other widespread taxa (sometimes having cosmopolitan or wide distributions all over the Mediterranean basin, occurring also in areas of Anatolia and the Balkan peninsula); (ENS): species of narrow distributions in the Aegean Sea, or endemics to a group of Aegean islands or even to a single island (Appendix). For the investigation of the faunal relationships within and between island groups, two slightly different strategies were followed. Firstly, four similarity coefficients were tested, two symmetrical (Pearson’s W and Yule) and two asymmetrical (Jaccard and Kulczynski #2), on a binary matrix of the data, using the NTSYS software (see Rohlf 1993 for equations). Since each group consists of one large island (the species ‘‘pool’’) and several very small satellites, symmetrical similarity coefficients (which consider equally – or via a more sophisticated estimation – the ‘‘zero’’ and the ‘‘1’’ states) are believed to better resolve the overall similarities. The great number of species absences from the small satellite islands provides useful biological information. Therefore, we favour Pearson’s W and Yule’s indices, whose values were used as input for UPGMA cluster analysis. Secondly, dissimilar weight was given on endemics plus subendemics (53), Balkan and Anatolian (52), and widespread (51) species on the above (binary) matrix due to the fact that the various species distributional categories provide different biogeographical information, and quantitative coefficients were applied to recalculate the similarities between islands. Four of the commonest quantitative similarity indexes were used: Bray-Curtis, Morisita, Pearson product-moment and Cosine (see Sneath
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496 A. Trichas et al.
Figure 4. Map of the island group of Nisyros, its relative position in the Aegean and the sampling stations (dots5hand collections, crosses5pitfall trapping stations).
and Sokal 1973). The clustering method was UPGMA again. Cophenetic correlation coefficients were used for the evaluation of the trees produced by both binary and non-binary matrixes (Rohlf 1993). Despite the plethora of proposed models to describe the species-area relationship, only the most commonly used proposed by Arrhenius (1921) was employed, mainly due to the fact that it is the only model so far, with parameters assigned having biological meaning (see Connor and McCoy 1979; Brown and Lomolino 1998). As usual, the model was applied in its logarithmic form: logs5logc+zlogA, where S is the number of species and A area in km2. For the exploration of nestedness patterns the BINMATNEST software was used (Rodrı´gue´z-Girone´s and Santamarı´a 2006). Higher temperatures are indicative of lower community nestedness, while ‘‘colder’’ matrices are more nested. The software developed by the authors calculates a ‘‘Temperature’’ value (T) for each community that is similar to the widely applied Nestedness Temperature Calculator (Atmar and Patterson 1993). Although the ‘‘Nestedness Temperature Calculator’’ method has been widely used in the literature and has become a familiar tool for many researchers (e.g. Sfenthourakis et al. 1999, 2004), it has some major limitations (Wright et al. 1998; Fischer and Lindenmayer 2002; Rodrı´gue´z-Girone´s and Santamarı´a, 2006). Thus, the BINMATNEST method was applied; it is based on the same principles as the ‘‘Nestedness Temperature Calculator’’ but overcomes some of the major shortcomings of the method (for analytical discussion see Rodrı´gue´z-Girone´s and Santamarı´a (2006)).
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For regression analyses STATISTICA (ver. 5.00 StatSoft, Inc. 1997) and SPSS (ver. 14.0, SPSS Inc. 1989–2005) were used. Results General observations and faunal remarks A total of 43 tenebrionid taxa was recorded, raising the number of known taxa from the area to 59 (44 were previously known, almost only from the large islands, with the majority – 32 – from Santorini island). Specifically, 19 taxa were recorded from Thira (5Santorini main island) and Thirasia, 11 from the Astypalaia island group, 17 from the Kalymnos group and 19 from the Nisyros group. So far, the specimens from Thira and Thirasia did not add any new records, a fact that reveals the thoroughness of the tenebrionid knowledge for these islands (Appendix). As, to present knowledge, no previous researchers have used pitfall traps for collecting tenebrionid beetles in the Aegean, these data provide useful semi-quantitative information on species abundances (as approximated by epedaphic mobility): through all the islands of interest, the most abundant genera (with hundreds of individuals per species) proved to be Dailognatha Eschscholtz, 1829, Dendarus Latreille, 1829, Graecopachys Skopin, 1968, Opatroides Brulle´, 1832, Pachyscelis Solier, 1836, Pimelia Fabricius, 1775, Raiboscelis Allard, 1876, Tentyria Latreille, 1802, and Zophosis Latreille, 1807. In some cases Dichomma Solier, 1835, Erodius Fabricius, 1775, Eutagenia Reitter, 1886, Pedinus Latreille, 1796, Sclerum Dejean, 1834 and Stenosis Herbst, 1799, were also abundant. Except for Dichomma, Opatroides and Sclerum, all other genera speciate intensively in the area, being responsible for the major part of Aegean endemicity (the most extreme example being the genus Dendarus – Chatzimanolis et al. (2002, 2003)). In one case (Astypalaia and almost all of the adjacent islets), dozens of individuals of Graecopachys quadricollis (Brulle´, 1832) were found, a species not previously recorded from the area, while Pachyscelis villosa (Drapiez, 1820), a species of similar appearance, was recorded by Lo Cascio and Scupola (2003) from the main island of Astypalaia, but this record was not confirmed. Nevertheless, both species are included in the analyses. Other points of interest were the species of the genus Dendarus. Except for Dendarus wernerianus Koch, 1948 from Santorini, all Dendarus taxa found in this study are new records for the area, with the most interesting being Dendarus anaphianus Koch, 1948 from Astypalaia and the adjacent islets. The species was described from the nearby island of Anaphi and was confirmed also from its neighbouring islets of Megalo Fteno, Mikro Fteno, Pacheia and Anydros by Chatzimanolis et al. (2002). It belongs to the sinuatus group of species (Koch 1948), a clade of Dendarus that is typically Aegean (Chatzimanolis et al. 2003; Trichas, unpublished data) and together with the Graecopachys quadricollis (also with Aegean affinities) from the same island, set Astypalaia at the edges of the Pleistocene Cycladic subcontinent. On the contrary, D. stampalicus Koch, 1948 (Stampalia5Astypalaia) of the rhodius species group (Koch 1948), reconfirmed by Chatzimanolis et al. (2002) on almost all small islets south of Astypalaia (Syrna, Zafrana, Astakida, etc.), was not recorded anywhere within the Astypalaia archipelago in the current study. Five different Dendarus species were found (four on Kalymnos island group and one on Nisyros, see Appendix). One species on Nisyros and a second one on Kalymnos seem to be of the D. rhodius Baudi, 1876 lineage; therefore they were
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498 A. Trichas et al. denoted as Dendarus cf. rhodius 1 and 2, respectively, for the purposes of the present analysis. A very small species from Kalymnos bears a close resemblance to D. tenellus (Mulsant & Rey, 1854), and is therefore referred to as D. cf. tenellus, and another one seems close to D. moesiacus (Mulsant & Rey, 1854) and is referred as D. cf. moesiacus. The fourth species from Kalymnos has no resemblance to any known Aegean form of Dendarus (and is probably an Anatolian taxon) and herein is referred to as D. sp1. The above species, as well as a new Stenosis, a new Catomus Allard, 1876 and a new Probaticus Seidlitz, 1891 form (Appendix) will be described in a forthcoming publication (Trichas A. in preparation). Chorological analysis The percentage of species in each chorological category is presented in Table 1. There is a clear cline of Anatolian affinities with the minimum percentage on Santorini (9%) and the maximum on Kalymnos (33%). Endemic plus subendemic taxa comprise almost the same proportion on all islands, except for Nisyros, which also lacks Balkan elements (see also Lo Cascio and Scupola 2003). Finally, taxa with wider distributions (occurring all over the eastern Mediterranean region) clearly dominate, with percentages ranking between 33% and 63%, the maximum recorded on Nisyros. Cluster analyses The UPGMA trees produced by both the asymmetrical (Jaccard and Kulczynski #2, cophenetic correlation coefficients: 0.91 and 0.90, respectively) and symmetrical (Pearson’s W and Yule, cophenetic correlation coefficients: 0.93 and 0.81, respectively) coefficients were very similar, with almost identical topologies, at least for the rough island group clustering. All similarity coefficients agreed in clustering the Santorini and Astypalaia groups together, apart from the eastern Aegean aggregates of Nisyros and Kalymnos, which in turn are fused together into a larger cluster. Pearson’s W coefficient gave the highest cophenetic correlation coefficient (0.93), so the respective dendrogram is given here (Figure 5). Similar topologies (but not identical) were produced by the transformed binary matrix (with dissimilar weights on species presences, as described previously). The four quantitative indices used, resulted also to similar dendrograms, with high cophenetic correlation coefficients (.0.9). The Cosine coefficient (with the highest cophenetic correlation50.96) was chosen for presentation of the results (Figure 6). Table 1. Chorological categories in the studied area (ANA5Anatolian, BAL5Balkan, ENS5Endemic plus Subendemic, WID5Widespread), based on the categories proposed by Fattorini et al. (1999). See text for further explanations. Island groups
ANA
BAL
ENS
WID
SANTORINI ASTYPALAIA KALYMNOS NISYROS
9% 16% 33% 29%
6% 11% 10% 0%
28% 16% 24% 8%
57% 57% 33% 63%
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Figure 5. The UPGMA tree based on the results of Pearson’s W coefficient applied on binary data. The prefixes SA, AS, KA and NI on the names of the islands, indicate the island group they belong (SA, Santorini group; AS, Astypalaia group; KA, Kalymnos group; NI: Nisyros group). The underlined names represent the main islands in each group.
Figure 6. The UPGMA tree based on the results of Cosine coefficient applied on the transformed, non-binary data matrix. The prefixes SA, AS, KA and NI on the names of the islands, indicate the island group they belong (SA, Santorini group; AS, Astypalaia group; KA, Kalymnos group; NI: Nisyros group). The underlined names represent the main islands in each group.
500 A. Trichas et al.
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Table 2. Results of the species-area analysis in the four island groups (per island group and the entire studied area). r, correlation coefficient. Island group
R
P
SANTORINI
0.961
50.039
z50.235 R250.923 c510.03
ASTYPALAIA
0.914
,0.001
z50.277 R250.835 c55.41
KALYMNOS
0.908
50.002
z50.248 R250.824 c55.21
All islands
0.867
,0.001
z50.278 R250.751 c56.04
Santorini and Astypalaia groups appear with similar topologies as above, but giving more importance to more localised taxa, Kalymnos and Nisyros groups also tend to cluster apart, showing a higher degree of geographical consistency (Figure 6). Species-area curves Species richness was significantly correlated with area for all island groups, except for the Nisyros archipelago, where the linear regression was not statistically significant, probably because of the small number of islands included. The values of the z and c parameters for each of the island groups, as well as for the total of islands studied, are given in Table 2. Nestedness Nestedness was detected in all four island groups. The remote groups of Astypalaia and Santorini are more nested than Kalymnos, while Nisyros is the least nested, with an ‘‘extreme’’ value of 14.25 (Table 3). Table 3. Nestedness analysis in the studied area. T, Nestedness values. Island group
Astypalaia Santorini Kalymnos Nisyros
BINMATNEST T
P
4.71 5.27 6.07 14.25
,0.001 ,0.001 ,0.001 ,0.001
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Discussion and conclusions Downloaded By: [University of Oxford] At: 14:19 3 June 2008
Chorological analysis and faunistics Tenebrionid beetles can solve many biogeographical issues in the Aegean area. They incorporate genera highly diversified in this area (i.e. Dendarus), and given their extremely limited dispersal abilities through aptery (Fattorini and Fowles 2005), each of them is useful for answering very particular biogeographic questions; sometimes at a ‘‘resolution’’ level of a single island. As earlier pointed out (Fattorini et al. 1999), the discovery rate of tenebrionid species in the Aegean archipelago has not yet reached a plateau. Moreover, new sampling methods (i.e. pitfall trapping) tend to add new species records on islands quite well investigated in the past, as indicated for some abundant genera (Dendarus, Graecopachys, etc.) in the present study. The addition of several tenebrionid beetle species after a more thorough sampling effort provided more reliable biogeographical patterns. The biogeographical profile of each island group reflects exactly its relative position in the Aegean and the degree of its remoteness or isolation, both geologically and geographically. The cline in Anatolian affinities (Table 1) clearly reflects the geography of the area. The somewhat lower representation of Anatolian elements on Nisyros, in comparison to Kalymnos, can be attributed to the combination of the relatively larger distance of the Nisyros group from the Anatolian coastline (see Figures 3 and 4), and to the difference in age and geological character of the two island groups (as already stated, Nisyros is a recent volcano). The same rationale explains also the poor representation of endemic plus subendemic elements on Nisyros and the absence of Balkan affinities on the same island. Kalymnos is a continental archipelago and, although recently isolated from the Anatolian coast (Van Andel and Shackleton 1982; Lambeck 1996; Perissoratis and Conispoliatis 2003), it always had its own established fauna with an evident dominance of Anatolian elements. The biogeographical position of Astypalaia and its satellite islets at the edges of the Cycladic and the Dodecanese provinces has been debated in the past, with evidence supporting both Cycladic and Dodecanese affinities for this island group (Sfenthourakis 1996a; Simaiakis 2006, and references therein). In Aegean tenebrionid studies, the naming of a Dendarus taxon distributed on several islets south of Astypalaia as D. stampalicus by Koch (1948) could be misleading as far as the biogeographical position of Astypalaia is concerned. The discovery of the true Astypalaian Dendarus (D. anaphianus) on the main island, as well as on the surrounding islets, given the importance of this speciose genus in the Aegean (Chatzimanolis et al. 2003), strongly supports the Cycladic character of the Astypalaia group, setting once more the south and south-eastern biogeographical limit of the Cycladic plate, exactly on Astypalaia island. Dailognatha quadricollis anaphiana Koch, 1948 and Graecopachys quadricollis also suggest the affinities of Astypalaia to be with Anaphi and other Cycladic islands. Cluster analyses All the UPGMA trees (resulted from both binary and non-binary data matrices) agreed in the identification of clearly separate Santorini and Astypalaia groups that are connected to each other, apart from the eastern Aegean (Kalymnos-Nisyros) cluster, accentuating the unique faunal character of the islands and the west-east
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502 A. Trichas et al. Aegean discontinuity. The Kalymnos-Nisyros groups constitute a separate cluster of islands and islets mixed together in various positions, depending on the similarity index used (in the presence-absence analysis). The use of Yule’s index in the same analysis failed to clearly separate the Kalymnos and Nisyros groups, but when localised taxa were weighted in the data matrix (leading to a non-binary transformation of the matrix), almost complete separation of the two groups (except for two Nisyros islets) was achieved. Moreover, there were no significant differences among indices in the patterns of isolation of the remote island groups of Santorini and Astypalaia. This, in concordance with the results of the nestedness analysis (see below), can be considered indicative of the balanced, ‘‘relaxed’’ and clearly defined character of the tenebrionid fauna of those islands. The fauna of Santorini and Astypalaia, islands clearly isolated from any nearby coast or larger landmass for several thousands of years, probably consists of the preexisting species (largely the subendemics and a few of the east Mediterranean taxa in the analysis) and, only secondarily, by newcomers (several of the widespread taxa in the analysis). According to Dermitzakis and Papanikolaou (1981) and Dermitzakis (1990), today’s parts of the Astypalaia group of islands, are remnants of Cycladic landmasses dating as old as the lower Miocene, which is in agreement with the relictual and balanced character of the established fauna on this island, indicating also probable ancestors of its endemic or subendemic taxa. Kalymnos and Nisyros are somewhat different cases and must be treated separately, but their proximity to the Anatolian coast and the small distance between them can satisfactory explain their relative positions in the cluster analyses. Finally, since the distances between almost all the satellite islets within each of the four island groups are very small compared to the whole studied area, and given the relatively small number of taxa on most of the islets, no further discussion on the particular position of an islet within a group can be made. Species-area analysis Strong and highly significant correlation was revealed between species richness and area, for all the island groups as well as for all the islands studied, with the exception of the Nisyros group. The same has also been stressed for many larger Aegean islands for tenebrionid beetles (Fattorini 2002; Fattorini and Fowles 2005), terrestrial isopods (Sfenthourakis 1996b), centipedes (Simaiakis 2006) and land snails (Mylonas 1982; Triantis et al. 2005a, 2005b; Triantis 2006). The slopes of the species-area relationship in each island group, as well as the overall slope, place the island groups in the archipelagic category but are quite close to the values reported for the within biogeographic province category (intraprovincial) as proposed by Rosenzweig (1995). Similar z values are found when we compare the diversities of different-sized islands. The relatively low z values for all of the island groups are the presumed result of the ‘‘recent’’ formation of the island groups (Van Andel and Shackleton 1982; Dermitzakis 1990; Lambeck 1996; Perissoratis and Conispoliatis 2003). A significant number of island groups in the Aegean have been formed quite recently in geological time, and even for taxa with reduced dispersal ability, the islands still ‘‘behave’’ as parts of a continuous land mass, with high numbers of species even on the smallest islands, marginal extinctions, limited net effects of island size and a significant contribution of the
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Journal of Natural History
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interrelationship between area and environmental heterogeneity. These result in relatively low slope values (see also Triantis et al. 2005a, 2005b; Triantis, 2006). The c value of all islands in the present study is 6.039, with 5.214,c,6.317 for the separate Kalymnos, Nisyros and Astypalaia archipelagos, and an exceptional c510.030 for the Santorini island group. The value given in the earlier study of Fattorini (2002) was 3.45, indicating an underestimation of the richness of the tenebrionid beetle fauna in the area. Applying a Student’s test (see Zar 1984) in order to compare the two respective c-values, a statistically significant difference was found between the two values (P(0.05). Similar studies in the Aegean archipelago with isopods and snails have given c values of 9.33 and 9.48, respectively (Sfenthourakis 1996b; Welter-Schultes and Williams 1999; Triantis 2006), which are much closer to the value of the Santorini archipelago for tenebrionid beetles. In our opinion, Santorini has been almost exhaustively sampled, as no new species were added after the present research, in contrary to all other groups of islands. Thus, a more realistic estimation for the c value of the Aegean tenebrionid beetles should be closer to the value of the Santorini group, with values of the other three groups being minimal ones. Therefore, for Aegean tenebrionid beetles the value of c should lie between 6 and 10. Using values of c510.03 and z50.235 for the largest Aegean island (Crete), one should obtain an estimation of 84 species. This value is quite close to the present number of species reported (c. 80; considering data from Koch (1948), Ku¨hnelt (1965), Kaszab (1973), Trichas (1996), and Fattorini (2006) and references therein). For the island of Santorini, there would be 28 species, with the actual number of species reported to be 32 (see Appendix). Nestedness According to BINMATNEST method all data matrices of the four island groups show significant nestedness. The ‘‘temperatures’’ calculated for each group fit quite well to the first assumptions on the biogeographical character of the archipelagos. The remote ones (Santorini and Astypalaia) are the most nested, with temperature values marginally different to each other. Bearing in mind the formation of these island groups and their degree of isolation in space and time (see also the discussion in Cluster analyses), it is obvious that for thousands of years the main islands of these groups were the only source of faunal exchanges with their satellite islands, hence increased nestedness was predictable. Higher, but significantly different between each other, were the temperatures of the coastal island aggregates of Nisyros and Kalymnos. Assuming their relative positions near the Anatolian coast, one can expect similar temperatures. But the different geological formation for these island groups has deeply affected their tenebrionid fauna. The isolation of the Kalymnos island group from the Asia Minor coasts is no older than 20,000 years, with some parts being connected to an elongated landmass including the islands of Leros and Arkoi before 11,000–9000 years (Van Andel and Shackleton 1982; Lambeck 1996; Perissoratis and Conispoliatis 2003). Today’s geography can be considered only 2000 years old (Papageorgiou 2001), but the clearly continental formation of the area (with established, balanced fauna) has not permitted major stochastic recolonizations of the islets by coastal newcomers. The tenebrionid taxa on the satellite islets of Kalymnos are subsets of the former
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504 A. Trichas et al. single landmass, showing a considerable degree of nestedness. On the other hand, the Nisyros group shared no permanent land bridges with the Anatolian coast and has a clearly oceanic character (not only the main island of Nisyros, but most of its satellite islets are of volcanic origin), and a much more stochastic faunal formation than Kalymnos. All taxa inhabiting the Nisyros group of islands are more or less recent immigrants from the nearby Anatolian coast or from other Dodecanese islands. The results of the chorological analysis (Table 1) are also in agreement with the above rationale: Nisyros hosts almost double the widespread taxa of the nearby Kalymnos and three times fewer endemic plus subendemic taxa. Finally, we would like to underline the importance of islet research in the Aegean area. There are several recent papers (Sfenthourakis 1996a, 1996b; Foufopoulos and Ives 1999; Dennis et al. 2000; Fattorini 2002; Chatzimanolis et al. 2003; Fattorini and Fowles 2005; Hausdorf and Henning 2005) dealing with Aegean biogeography and most of them agree on the basic distribution patterns in this field. There are no doubts so far about the unique biogeographic position of Crete and the south Aegean arc, the east–west Aegean discontinuity or other similar large scale patterns (Sfenthourakis 1996a; Hausdorf and Henning 2005, and references therein). It is the fine details in the patterns that have to be revealed in the proximate future and focusing on smaller archipelagos, choosing them properly and using appropriate tools, will gradually lead to a complete scenario for the biogeography and evolution in the Aegean area. Acknowledgements We wish to thank Dr P. Lyberakis, D. Kaltsas, M. Nikolakakis and K. Madi for their valuable help during field trips and the sorting of the coleopterological material, and Dr A. Parmakelis who kindly made many valuable comments on an early version of the manuscript. The present study was included in the program PYTHAGORAS I ‘‘Multilevel approach of biodiversity on four island groups in the east Aegean’’ and co-funded by the European Union (75%) and by the Operational program EPEAEK II (Education and Initial Vocational Training) of the Ministry of National Education and Religious Affairs.
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8 9 10 11 12 13 14 15
WID ANA – BAL ENS
Nera Sari Nisyros Gyali Pacheia Kandeliousa Pergousa
Kalavro
Plath Agia Kuriakh
Pserimos
Telendos
Kalymnos
Fteno
Megalo Fokionisi
Mikro Fokionisi
X
Pontikousa
– ANA
Ofidousa
X
Koutsomytis
WID
Agia Kyriaki
X
Kounoupi
L
Astypalaia
WID BAL WID WID ENS
Aspro
X X
Pal. Kameni
WID WID
Thirasia
Santorini
3 4 5 6 7
Akis elongata Brulle´, 1832 Alphitobius diaperinus (Panzer, 1797) Ammobius rufus Lucas, 1849 Belopus reitteri (Zoufal, 1893) Blaps mucronata Latreille, 1809 Blaps tibialis Reiche, 1857 Cataphronetis reitteri (Seidlitz, 1894) Catomus consentaneus (Ku¨ster, 1851) Catomus sp.1 Cephalostenus orbicollis Me´ne´trie´s, 1836 Cheirodes sardous Ge´ne´,1839 Colpotus sulcatus (Me´ne´trie´s, 1838) Crypticus sp. Dailognatha hellenica Reitter, 1896 Dailognatha quadricollis anaphiana Koch, 1948
DISTRIBUTION
1 2
AEGEAN
Taxa
L
X
L
L L L N
L
N
N X
N N N N
L L
L
X
L
N L
N X
L
N
L N
X
N
N
L X
N
N
N
X
N N N N
X
508 A. Trichas et al.
Appendix. List of tenebrionid species in the island groups. X: presence, N: New reference, L: Literature data only. For the Aegean distributions refer to text.
ENS ENS ENS ENS ENS ENS ENS ANA
N X
X
N
N
N
N
Nera Sari Nisyros Gyali Pacheia Kandeliousa Pergousa
Kalavro
Plath Agia Kuriakh
Pserimos
Telendos
Fteno
N
L N N N N N N N N
X L
ANA ENS
X
X
ANA WID
X
X
L
L
X L L
WID WID
Kalymnos
Megalo Fokionisi
Mikro Fokionisi
Pontikousa
Ofidousa
Koutsomytis
Agia Kyriaki
Kounoupi
Astypalaia
Aspro
Pal. Kameni
Thirasia
DISTRIBUTION
Santorini
X N X N N N N N X N N N N
ENS
L
L
L
N
N
N L
N
X N N
N
X N N L
Journal of Natural History 509
16 Dailognatha quadricollis sporadica Koch, 1948 17 Dendarus anaphianus Koch, 1948 18 Dendarus wernerianus Koch, 1948 19 Dendarus sp1 20 Dendarus cf moesiacus 21 Dendarus cf rhodius 1 22 Dendarus cf rhodius 2 23 Dendarus cf tenellus 24 Dichomma dardanum (Steven, 1829) 25 Erodius orientalis oblongus Solier, 1834 26 Erodius orientalis brevicostatus Solier, 1834 27 Eutagenia minutissima Pic, 1903 28 Eutagenia smyrnensis (Solier, 1834) 29 Gonocephalum affine (Billberg, 1815) 30 Gonocephalum rusticum (Olivier, 1811)
AEGEAN
Taxa
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Appendix. Continued.
WID WID WID ANA BAL WID ENS WID
L
L
L
X
L
L
X
N
X N N
L N L
X X
X N N
Nera Sari Nisyros Gyali Pacheia Kandeliousa Pergousa
Kalavro
Plath Agia Kuriakh
Pserimos
X
N
Telendos
WID
N
Fteno
L
N
Kalymnos
L L
N
Megalo Fokionisi
ENS ENS
N
Mikro Fokionisi
L
Pontikousa
X
Ofidousa
X
Koutsomytis
ENS
Agia Kyriaki
L
Kounoupi
L
Astypalaia
Thirasia
L
Aspro
Santorini
WID
Pal. Kameni
DISTRIBUTION
31 Gonocephalum setulosum (Faldermann, 1837) 32 Graecopachys quadricollis (Brulle´, 1832) 33 Gunarus kaszabi Grimm, 1981 34 Micrositus orbicularis Mulsant & Rey, 1854 35 Opatroides punctulatus Brulle´, 1832 36 Opatrum geminatum Brulle´, 1832 37 Opatrum obesum Olivier, 1811 38 Opatrum sabulosum (L., 1758) 39 Pachyscelis villosa (Drapiez, 1820) 40 Pedinus quadratus Brulle´, 1832 41 Phaleria bimaculata (Linne´, 1767) s.l. 42 Pimelia sericella Solier, 1836 43 Pimelia subglobosa (Pallas, 1781) s.l. 44 Polycoelogastridium sexcostatum (Motschulsky, 1858) 45 Probaticus sp1
AEGEAN
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Taxa
N N
N X
N
N N
L L N L X N
N
N L
X L
WID
N
–
N
N
N N
N N
N
510 A. Trichas et al.
Appendix. Continued.
Nera Sari Nisyros Gyali Pacheia Kandeliousa Pergousa
Kalavro
Plath Agia Kuriakh
N N N
Pserimos
N
Telendos
N
Fteno
Pontikousa
N
Kalymnos
Ofidousa
N
Megalo Fokionisi
Koutsomytis
X
Mikro Fokionisi
Agia Kyriaki
L
Kounoupi
Pal. Kameni
L
Astypalaia
Thirasia
L
Aspro
Santorini
46 Raiboscelis azureus obliteratus BAL Allard, 1878 47 Raiboscelis coelestinus ANA (Waltl, 1838) s.l. 48 Sclerum multistriatum WID Dejean, 1834 49 Stenosis esau Sah.lberg, 1907 ANA 50 Stenosis syrensis Koch, 1936 ENS 51 Stenosis sp.1 – 52 Tentyria rotundata mittrei ANA Solier, 1835 53 Tentyria rotundata sulcatipennis ENS Schuster, 1915 54 Trachyderma lima (Petagna, WID 1819) 55 Trachyscelis aphodioides WID Latreille, 1809 WID 56 Tribolium castaneum (Herbst, 1797) 57 Tribolium confusum Jaqcquelin WID du Val, 1868 58 Zophosis dilatata Deyrolle, 1867 WID 59 Zophosis punctata Brulle´, 1832 WID
Taxa
AEGEAN
DISTRIBUTION
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Appendix. Continued.
X N X N N N X
L
N N
N
X
N N N
N
N N N
N X
N
L X L X
X
L
N
L
L
L
L L
X
X
L
L N N
N N
X X N
N
Journal of Natural History
X
511
