Anterastes Brunner von Wattenwyl, 1882

Uluar, Onur, Yahyaoğlu, Özgül, Başıbüyük, Hasan Hüseyin & Çıplak, Battal, 2023, Taxonomy of the rear-edge populations: the case of genus Anterastes (Orthoptera, Tettigoniidae), Organisms Diversity & Evolution (New York, N. Y.) 23 (3), pp. 555-575 : 567-572

Genus Anterastes Brunner von Wattenwyl, 1882

1- A. disparalatus Ünal, 2002

2- A. antecessor Kaya & Çıplak, 2011

A. uludaghensis species group

3- A. uludaghensis Karabağ, 1950

4- A. davrazensis Kaya et al., 2015

A. babadaghi species group

5- A. niger Ünal, 2000

6- A. turcicus Karabağ, 1951

7- A. babadaghi Uvarov, 1939

8- A. ucari Çıplak, 2004

9- A. kizkayasiensis Çıplak & Uluar sp.n

A. serbicus species group

10- A. serbicus Brunner von Wattenwyl, 1882

11- A. boreoanatoliensis Çıplak & Uluar sp.n

12- A. burri Karabağ, 1951

13- A. muratdagensis Çıplak & Uluar sp.n

14- A. tolunayi Karabağ, 1951

15- A. antitauricus Çıplak, 2004

16- A. anatolicus Uvarov, 1934 (known only from the type material and fresh material was not available to include in this study)

A. kizkayasiensis Çıplak & Uluar sp.n.

LSID urn:lsid:zoobank.org:act:BF085BEB-AFE3-4965–9752-07B12DB3D0C6 .

Description: See Appendix 1.

A. boreoanatoliensis Çıplak & Uluar sp.n.

LSID urn:lsid:zoobank.org:act:98C005E3-DDED-4C45-B51F-BE4122C4890E .

Description: See Appendix 1.

A. muratdagensis Çıplak & Uluar sp.n.

LSID urn:lsid:zoobank.org:act:682441CF-FFDF-48BC-B32D-629070BFA231 .

Description: See Appendix 1.

Conclusions: taxonomy as a reflection of the glacial ages

We attempt to provide a reconciliation for the taxonomy of Anterastes + Koroglus using a data-based approach without subjective personal decision. Ten species in the genus restricted to a particular summit indicating their cold preferring habitat preference (Çıplak, 2004; Çıplak et al., 2010, 2015). Of these ten species, A. disparalatus , A. antecessor , A. uludaghensis , and A. davrazensis constitute the basal branches of the genus and ultimate ancestor of each date to Plio-Pleistocene transition or earlier. A. serbicus is also an old species by age of its ultimate ancestor dated to the Plio-Pleistocene transition. Although their ultimate ancestors are younger than previous species by the age of> 2.0 myr in the Early Pleistocene, A. burri , A. turcicus , A. tolunayi , A. niger , and A. muratdagensis sp. n. also exhibits a similar pattern for distribution and habitat preference, and also can be related to glacial cycles. There are further interesting aspects in their distribution pattern. For example, A. tolunayi and A. muratdagensis sp. n. are sister groups occupying two adjacent highlands. However, distributions of sister group pairs, namely A. uludaghensis / A. davrazensis and A. turcicus / A. niger are separated by almost 1000 km without any records in between. Remaining species A. antitauricus , A. boreoanatoliensis sp. n., A. babadaghi , A. ucari , and A. kizkayasiensis sp. n. are the younger species by ultimate ancestor with upper bound as MPT. Of these, A. kizkayasiensis sp.n. possibly established by a marginal population split from ancestral stock shared with A. ucari as a possible bud species (Kaya & Çıplak, 2016). Other four young species A. antitauricus , A. boreoanatoliensis sp. n., A. babadaghi , and A. ucari possibly experienced subsequent range expansion/ contractions more than once after MPT or during last four glacial periods. Each scattered population to adjacent highlands is a likely remnant of these subsequent range changes.

The above assertions may allow us to accomplish some generalizations and offer some conceptual reasoning on the speciation pattern of glacial rare edge populations. First, all members of the genus prefer cold climates and are mountainous species, and this habitat preference seems to be conserved throughout the evolutionary history of the genus. Second, the range change patterns of such populations occur as an expansion to larger ranges in lowlands/southward during cold periods and the opposite during interglacial periods. Ten species restricted to single summit/chains and patchy populations of the widespread species are representative of the remnants during these range shifts. The old remnants before MPT constitute present species and those after the MPT present patch or trailing edge subpopulations of a widespread species. Thus, we assume the relicts before MPT as the stable rear-edge populations (Hampe & Petit, 2005). Furthermore, five species with relatively wider ranges represent the ancestral stock managed to expand their ranges during recent glacial cycles, while those restricted to a single highland/highland chain did not expand their range and remained relicts. Rapid divergence of the small population is a common evolutionary trend. In general, most of Anterastes species and specifically the species with an ultimate ancestor within Early Pleistocene, namely A. uludaghensis , A. davrazensis , A. turcicus , A. niger , A. kızkayasiensis sp. n., A. muratdagensis sp. n., A. tolunayi , and A. burri experienced rapid speciation events originated from such relict ancestral stocks. However, due to conserved habitat preference or tracking similar ecological conditions, they were subjected to similar selection pressures and slightly diverged in morphology (Ackerly, 2009; Cadotte et al., 2013), as revealed by the high phylogenetic signal in binary and metric characters. Taking all evidence together indicate that MPT or transition of cold periods from 42 to 100 kyr constitute a prominent stage to be considered in species level taxonomical decision for the stable rear-edge populations, if an age is going to be assigned to a species taxon (Avise & Liu, 2011). We highlight a new issue to be investigated by further studies in lineages with similar habitat preferences and phylogenetic history. In this context, it is interesting to note that a recent study (Ehl et al., 2021) obtained similar results for indicating a likely generality of the pattern.

Appendix 1. Descriptions of three new species given in the study

Genus Anterastes (Brunner von Wattenwyl, 1882) tic structures (female subgenital plate and ovipositor) of sister species Anterastes ucari are given in the box for the comparison.] Anterastes kizkayasiensis Çıplak & Uluar sp.n.

( Fig. 5 ).

LSID urn:lsid:zoobank.org:act:BF085BEB-AFE3-4965–9752-07B12DB3D0C6 .

Holotype, ♂, TURKEY: Konya, Kızkayası Mt. , 38° 05′ 019.7″ N, 32°16′ 056.0″ E, 1644 m, 01.07.2008 (Çıplak et al.) ( MEVBIL) . Paratypes, 7♂, 9♀, same data as holotype .

Diagnosis: As stated by genetic data, the new species constitutes an independent phylogenetic and reproductive unit within A. babadaghi group (given as A. cf. ucari in Çıplak et al., 2015). It can be differentiated from other members of the group by the allopatric species range distribution. The new species share morphological features of A. babadaghi group (see Çıplak, 2004), but more similar to A. ucari and A. babadaghi . Although differences are not prominent, it is not a cryptic species and can be distinguished from these two by a smaller size and especially short, slender, and almost straight nostic structures (female subgenital plate and ovipositor) of sister species Anterastes serbicus are given in the box for the comparison.] ovipositor and narrower incision of the female subgenital plate. The PCA applied to 23 metric characters well distinguish it from other members of the species group ( Fig. 4 ). In addition to phenotype, the ML tree obtained from full data set subjected to PAUP analysis defined the following unique base changes for the new species; 248 (A-> G), 683 (T-> C), 692 (A-> C) and 1268 (A-> G). The lists of the unique and non-unique base changes is presented in Fig. 2 and File S1.

Etymology. The species was named after the type locality, the Kızkayası Mt. in Konya provinces of Turkey.

Anterastes boreoanatoliensis Çıplak & Uluar sp.n.

( Fig. 6 ).

LSID urn:lsid:zoobank.org:act:98C005E3-DDED-4C45-B51F-BE4122C4890E .

Holotype, ♂, TURKEY: Giresun, Şebinkarahisar , 40° 27′ 046.2″ N 38° 42′ 048.3″ E, 2294 m, 22.07.2005 (Çıplak et al.) ( MEVBIL).

Paratypes, 45♂, 40♀, same data as holotype ; 18♂, 29♀, Sivas, Kösedağ Mt. , 40° 09,029.7″ N, 37° 51′ 002.6″ E ,

tic structures [female subgenital plate and ovipositor) of sister species Anterastes burri are given in the box for the comparison.]

1802 m, 23.07.2009 (Çıplak et al.); 13♂, 5♀, Kastamonu, Ilgaz Mt. , 41° 03′ 024.3″ N, 33° 43′ 005.8″ E , 2045 m, 13.08.2004 (Çıplak et al.); 11♂, 29♀, Çankırı, Çerkes, Işık Mt. , 40° 41′ 027.8″ N 32° 44′ 006.4″ E , 1595 m, 22.06.08 (Çıplak); 2♂, Erzincan, Pöske Mt. , 39° 53′ 26.4′′ N, 39° 21′ 43.3′′ E , 2169 m, 19.07.2016 (Çıplak & Kaya) ( MEVBIL) .

Diagnosis: The new species exhibits morphological features of A. serbicus group (given as A. cf. serbicus in Çıplak et al., 2015). It is morphologically cryptic and cannot be diagnosed from other members of the group. Especially, it is similar to A. serbicus and A. antitauricus by the titillators without curved apical arms. However, genetic data suggest it as a wellsupported independent phylogenetic unit. It can be properly diagnosed from other members of the group by its allopatric range. Although this new species occurs as sympatric with A. niger in Sivas, each belongs to different species groups. The ML tree of full molecular data displayed following unique base changes for the new species; 21 (A-> G), 205 (A-> G), 626 (C-> T), 627 (C-> T) 653 (T-> A), 662 (T-> G), and 1160 (A-> T) (see Fig. 2 and File S1).

Etymology: Range of this new species covers western and northern Black Sea Basin or boreal part of Anatolia and the species name was established after this eco-geographic feature of the species’ range.

Anterastes muratdagensis Çıplak & Uluar sp.n.

( Fig. 7 ).

LSID urn:lsid:zoobank.org:act:682441CF-FFDF-48BC-B32D-629070BFA231 .

Holotype, ♂, TURKEY: Uşak, Murat Mt. , 38° 56′ 054.7″ N, 29° 37′ 029.4″ E, 1785 m, 17.07.2009 (Çıplak et al.) ( MEVBIL).

Paratypes, 16♂, 21♀, same data as holotype .

Diagnosis: The new species exhibits morphological features of A. burri especially by the apically bent apical arms of titillators, and also general characteristics of A. serbicus group (given as A. cf. burri in Çıplak et al., 2015). It can poorly be distinguished from other members of the group by pronotum (the length, width, and length/width of the pronotal disk in male and female). Genetic data suggest it as an independent phylogenetic and reproductive unit. Although it cannot be diagnosed by morphology, it can be well diagnosed from other members of the group by its range allopatric to all other members of the genus. The new species has the following unique base changes in the ML tree of full data set; 25 (A-> T), 284 (A-> G), 335 (G-> T), 569 (A-> T), 653 (T-> A), 770 (A-> C), 1076 (A-> C), 1157 (C-> T), 1394 (C-> T), and 1565 (T-> C) (see Fig. 2 and File S1).

Etymology: The species was named after the type locality of the species, the Murat Dağ Mt. in Uşak province in Aegean Basin of Turkey.

Figures

Fig. 5 Morphologic structures of Anterastes kizkayasiensis sp. n.: A pronotum+tegmina from above and profile, B male anal tergum and cerci, C titillators, D female subgenital plate, E ovipositor [diagnos-

Fig. 4 The PCA plots and trait contributions prepared from the metric data sets of A. serbicus (A) and A. babadaghi (B) species groups

Fig.2 BEAST chronogram obtained from MT and FD matrices with posterior probability supports (nodes received>0.9 posterior probability support in MT and FD chronograms were indicated by a bold circle.

Fig. 6 Morphologic structures of Anterastes boreoanatoliensis sp. n.: A pronotum+tegmina from above and profile, B male anal tergum and cerci, C titillators, D female subgenital plate, E ovipositor [diag-

Fig. 7 Morphologic structures of Anterastes muratdagensis sp. n.: A pronotum+tegmina from above and profile, B male anal tergum and cerci, C titillators, D female subgenital plate, E ovipositor (diagnos-