Endemic Carabus Species of Turkey: Some Researches with Biogeographic and Ecological Notes (Coleoptera, Carabidae, Carabinae)

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Endemic Carabus Species of Turkey: Some Researches

with Biogeographic and Ecological Notes

(Coleoptera, Carabidae, Carabinae)

Abstract

The endemic species of the genus Carabus occuring in Turkey are presented in this study. The data of the endemic species of Turkish Carabus recorded from our previous data collections, catalogues, and other various papers prepared on Carabinae, up until now were used. As a result, 35 endemic species and subspecies of Carabus in Turkey were recorded. The endemic species list of Turkish Carabus are presented together with distribution information. Additionally, the biotopes, density of species, and general information about whether they are officially under threat as a species is also presented. This information is very important for ecological and biological control, and for other such studies. Moreover, photos of some endemic Carabus are presented to be used in the diagnosis of these species. The aim of this study is to contribute a synthesis of information on the Turkish Carabus endemic species here in Turkey. Keywords: Biotopes, carabus, coleoptera, endemic species.

Türkiye’nin Endemik Carabus Türleri: Biyocoğrafik ve Ekolojik Notlarla Bazı Araştırmalar (Coleoptera, Carabidae, Carabinae)

Özet

Türkiye’de bulunan Carabus cinsine ait endemik türler bu çalışmada sunulmuştur. Türkiye’nin endemik Carabus türlerinin listesi bizim önceki toplama bilgilerimiz, Carabinae üzerine şuana kadar yayınlanan kataloglar ve değişik yayınlar kontrol edilerek hazırlanmıştır. Sonuç olarak, Carabus’un endemik 35 tür ve alttürü Türkiye’den kaydedilmiştir. Carabus’un endemik tür listesi türlerin yayılış bilgisi ile birlikte verilmiştir. Ayrıca türlerin biyotopları, yoğunluğu ve tehdit altındaki türlerin resmi olarak korunup korunmadığı ile ilgili genel bilgilerde sunulmuştur. Bu bilgiler ekolojik, biyolojik kontrol ve benzeri çalışmalar için çok önemlidir. Bazı endemik Carabus türlerinin fotoğrafı bu türlerin teşhisinde kullanılması için sunulmuştur. Bu çalışmanın amacı Türkiye’de yayılış gösteren endemik Carabus türlerinin bilgisine katkı sağlamaktır.

Anahtar Kelimeler: Biyotoplar, carabus, coleoptera, endemik türler.

Avgın SS, Prunier D (2015) Endemic Carabus Species of Turkey: Some Researches with Biogeographic and Ecological Notes (Coleoptera, Carabidae, Carabinae). Ekoloji 24(96): 26-40.

Received: 12.11.2014 / Accepted: 21.02.2015

Sakine Serap AVGIN1_{, Daniel PRUNIER}2

1_{Kahramanmaras Sutcu Imam University, Faculty of Education, Division of Science Education,} Avsar Campus, 46100, Kahramanmaras, TURKEY

2_{5 rue de l'Epargne, 92320, Chatillon, FRANCE}

*_{Corresponding author: ssavgin@ksu.edu.tr}

INTRODUCTION

Endemic species provide an important signal in conservation planning because they may indicate special qualities of the biotope in which they live that are not found elsewhere. Therefore, the first step in conservation planning is the preparation of lists of such species with notes about their way of life. Along with such lists, images of both habitats and the beetles provide tools for researchers and conservationists to further document why such taxa are indicators of areas to be protected. In the present study of the genus Carabus, additional information maybe acquired about their role in biological control since they are voracious predators.

Some species of Carabidae, like some species of

other insects are endemic. In various investigations, a checklist of the endemic Carabidae has been accumulated by different researchers (Gueorguiev 2007, Abdel-Dayem 2012). In Egypt, including the Sinai, there were 39 endemic carabid species and subspecies recorded and were arrayed across 28 genera by Abdel-Dayem (2012). The biogeography of endemic Carabidae belonging to 385 species and subspecies in the Central and eastern Balkan Peninsula was reported by Gueorguiev (2007).

There are endemic species belonging to different genera of Carabidae in Turkey. One of these genera is Carabus. The genus Carabus contains about 850 species in the world (Deuve 2006), and among those 337 species and subspecies are in Turkey (Avgın and

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Cavazzuti 2011).

Many studies have been made on Turkish Carabinae including Carabus by domestic and foreign researchers, but the vast majority of the studies have been conducted by researchers from abroad. In these studies, records of new species and distribution of species have been particularly presented. A contribution covering the endemic Turkish Carabus has not been offered until now.

Knowing the endemic species of Turkish Carabus

is important because many species of Carabus are predators and they can be used in biological control. But firstly, it is necessary to have information about these species and distributions of them.

Because of various human impacts over the next 30 years, or less, there will be many changes in nature, such as on the quality of the biotopes of species. Photographs of natural environments and species living there in is knowledge certain to be important for future research and conservation actions.

The list of Turkish endemic Carabus species, information on the distribution of these species, and the photos of them for the use in the diagnosis are presented in this study. Additionally, the quality of biotopes of these species and photographs of the natural environments of the Carabus species are also presented. The aim of this study is to contribute a synthesis of information on the Turkish Carabus

endemic species in Turkey.

MATERIAL AND METHODS

Endemic species of Turkish Carabus recorded from previous data collections, catalogues, and various papers prepared on Carabinae, up until now were used. This list was prepared especially using papers of Casale and Taglianti (1999), Turin et al. (2003), Smetana (2003), Deuve (2004), Cavazzuti (2006), Löbel and Avgın and Cavazzuti (2011). The photos of species and subspecies are presented together with their distribution in Turkey. The photos of many specimens were taken by Daniel Prunier. The quality of the biotopes are annotated as well as photographs of the natural environments of the species.The common species and general information about whether the species are under threat is summerized.

Size: Measurements of length (ABL) follow those of Ball (1972) and Kavanaugh (1979): ABL (apparent body length), measured from apex of labrum to apex of longer elytron (in adults of this

genus, the abdomen often protrudes beyond the elytral apex.

Type locality: The information was based on first collected region.

Material examined: The specimens examined in this paper are annotated. Photos of these specimens are presented.

Geographical range: The distribution of species in Turkey are given.

Chorotype: The distributional information is based on Taglianti et al. (1999).

Way of life and conservation: The quality of the biotopes of the species and photographs of the natural environments of the species are presented.

RESULTS

Endemic Species List of Turkish Carabus

Only photos of some species and subspecies and their biotopies are presently available and these are presented in this study. Result are given for 35 endemic species and subspecies belonging to the Turkish Carabus.

Family: Carabidae

Carabus Linné, 1758 (subg. Ischnocarabus

Kraatz, 1877)

Carabus (Ischnocarabus) tenuitarsis Kraatz, 1877

Distribution s.lat. (all ssp. known): Central and Northern Anatolia.

Type locality: N Turkey "Amasya" (Deuve 2004). Specimens examined: ssp. cankirianus Breuning 1962 from Cerkes 1100m, Çankırı (leg. I. Rapuzzi), ABL = 20/21mm.

Geographical range: Central and northern Anatolia (Deuve 2004).

Chorotype: This distribution exemplifies a Central Anatolian endemic chorotype.

Way of life and conservation: Lives on trays in wet pastures. Species affected by overgrazing.

Carabus (Ischnocarabus) cychropalpus Peyron, 1858

Distribution s.lat. (all ssp. known): Central and Southern Anatolia.

Type locality: S Turkey, Adana "Cydnus river"(Deuve 2004).

Specimens examined: MuseumNational d'histoire Naturelle (Paris), not available for the study.

Geographical range: Central and Southern Anatolia (Deuve 2004).

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Southern Anatolian endemic chorotype.

Way of life and conservation: This species has not recently been found. We do not know how it lives and this will require extensive research.

Carabus Linné, 1758 (subg. Tomocarabus

Reitter, 1896)

Carabus (Tomocarabus) simardianus Deuve, 1990

Distribution s.lat. (all ssp. known): Northern Anatolia near Amasia.

Type locality: N Turkey, near Amasya (Deuve, 2004).

Specimens examined: In private collection, not available for the study.

Geographical range: Northern Anatolia, near Amasya (Deuve 2004).

Way of life and conservation: Unknown.

Carabus (Pachystus) pisidicusPeyron, 1854

Distribution s.lat. (all ssp. known): Southern Anatolia, Taurus.

Type locality: SC Turkey, Antalya, Caraman (Deuve 2004).

Specimens examined: spp. pisidicusPeyron, 1854 from nord Ermenek, Konya, ABL= 25-28mm.

Geographical range: Southern Anatolia: Taurus (Deuve 2004).

Chorotype: This distribution exemplifies a Southern Anatolian endemic chorotype.

Way of life and conservation: Species of wide ecological amplitude that lives in the forest and maquis. Species of concern because of overgrazing in forests.

Carabus Linné, 1758 (subg.Oreocarabus

Géhin, 1876)

Carabus (Oreocarabus) porrectangulus Géhin, 1885

Distribution s.lat. (all ssp. known): Turkey, Pontic Alps: between Ordu and İspir, near Kastamonu and Bolu (Deuve 2004).

Type locality: N Turkey, Pontic Alps near Trabzon (Deuve 2004).

Specimens examined: Düzköy Yaylası, 1700 m, 1 specimen; Maçka-Sumela, 1 specimen, ABL = 24-27 mm.

Geographical range: Turkey, Pontic Alps: between Ordu and İspir, near Kastamonu and Bolu (Deuve 2004).

Chorotype: This distribution exemplifies a Northern Anatolian endemic chorotype.

Way of life and conservation: Species of wide ecological amplitude that lives in the forest and pasture altitude. Species of concern because of Fig. 1.Carabus(Lipaster) gordiusReitter, 1898: A-D; Habitus

in dorsal view (A: ssp. gordiusReitter 1898 from Kafkasor 1700m, Artvin, (Col. B.Lassalle), ABL = 36/38mm; B: ssp. bontempsiLassalle 1992 from Maçahel Gecidi, 1450m, Borçka, Artvin, (Col. D. Prunier), ABL = 35/40mm; C: ssp. osellaiBreuning & Ruspoli 1970 from Col de Cankurtaran, 800m, Hopa, Artvin (Col. B.Lassalle), ABL = 39/46mm; D: ssp. osellaiBreuning et Ruspoli 1970 from Hopa, Cankurtaran 800m; and E-F: biotope (E: from Hopa Cankurtaran 800m; F: from Hopa Cankurtaran Çiftepınar 520m).

Fig. 2.Carabus (Ischnocarabus) tenuitarsisKraatz, 1877, A: Habitus in dorsal view (A: ssp. cankirianusBreuning 1962 from Cerkes 1100m, Çankırı (leg. I. Rapuzzi), ABL = 20/21mm).

Fig. 3.Carabus (Pachystus) pisidicusPeyron, 1854, A: Habitus in dorsal view (A: spp. pisidicusPeyron, 1854 from nord Ermenek, Konya, ABL = 25-28mm).

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overgrazing in forests and pastures.

Carabus (Oreocarabus) mahouxiLedoux, 1972

Distribution s.lat. (all ssp. known): mountains south and southeast of Van Lake (Deuve 2004).

Type locality: SE Turkey, Van, 28 km N of Baskale (Deuve 2004).

Specimens examined: In private collection, not available for study.

Geographical range: East of Turkey, mountains south and southeast of Van Lake (Deuve 2004).

Chorotype: This distribution exemplifies an Eastern Anatolian endemic chorotype.

Way of life and conservation: Species of concerned because of overgrazing in forests and pastures.

Carabus Linné, 1758 (subg.Oxycarabus

Semenov, 1898)

Carabus (Oxycarabus) saphyrinus De

Cristoforis et Jan, 1837

Distribution s.lat. (all ssp. known): Turkey, Northern Anatolia.

Specimens examined: spp. saphyrinus De Cristoforis et Jan, 1837 from Kalecik 600m, Est Trabzon, 1 specimen, ABL = 27-32 mm; spp.

pseudosaphyrinus Schweiger, 1962 from Abant 1000m, Bolu, 1 specimen, ABL = 24-29mm; spp.

notabilis Roeschke, 1898 from Uludag, 1500m, Bursa, 1 specimen, ABL = 24-30mm; spp. pirithous

Reitter, 1896 from Ömerli, Alem Dağ, 1 specimen, ABL = 26-31mm.

Geographical range: Turkey, northern Anatolia (Deuve 2004).

Way of life and conservation: Species of wet forests and is not threatened by moderate logging. Fig. 4.Carabus (Oreocarabus) porrectangulusGéhin, 1885: A,

B: Habitus in dorsal view (A: spp., from Düzköy Yaysalı 1700m; B: spp., from Maçka-Sumela, Anatolie sept., ABL = 24-27mm).

Fig. 5.Carabus (Oxycarabus) saphyrinusDe Cristoforis et Jan, 1837: A, B, C, D: Habitus in dorsal view (A: spp. saphyrinusDe Cristoforis et Jan, 1837 from Kalecik 600m, Est Trabzon, ABL = 27-32 mm; B: spp. pseudosaphyrinusSchweiger, 1962 from Abant 1000m, Bolu, ABL = 24-29mm; C: spp. notabilisRoeschke, 1898 from Uludag, 1500m, Bursa, ABL = 24-30mm; D: spp. pirithousReitter, 1896 from Ömerli, Alem Dağ, ABL = 26-31mm).

Fig. 6.Carabus (Heterocarabus) mariettiCristofori et Jan, 1837: A, B, C, D, E, F, G, H: Habitus in dorsal view (A: ssp. sapancaensisBlumenthal & Breuning 1967

from Şenköy Teşvikiye, Yalova, ABL = 21/25mm; B: ssp. pseudomariettiSchweiger 1964 from Yeşigöller/Yığılca 1150m, Bolu, (leg. K.Staven), ABL = 17/22mm; C: ssp. necopitanusBasquin & Darge 1986 from İsmet paşa, ABL = 17/19mm; D: ssp. charetianusAuvray 1992 from Demirköy, 395/640m, Est Edirne, (Col.D.Prunier), ABL = 19/23mm; E: spp. bischoffiChaudoir, 1848 from Yenigun, 800m, Samsun, (Col. B.Lassalle), ABL =

18-22mm; F: spp. ornatusHeinz, 1976, from Yaraligöz (leg. Ph. Vingler), ABL = 17-20mm; G: spp. mucheiBreuning, 1961 from Ilgaz Geçidi, Kastamonu, ABL = 19-21mm; H: spp. akensis

Haury, 1889 from Ulukoy 1500m, Bursa, Uludag, ABL = 20-23mm) and I: biotope (I: from İğneada, Demirköy).

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Common.

Carabus Linné, 1758 (subg. Heterocarabus

Morawitz, 1886)

Carabus (Heterocarabus) marietti Cristofori et Jan, 1837

Distribution s.lat. (all ssp. known): Turkey, Northwestern Anatolia.

Specimens examined: ssp. sapancaensis

Blumenthal & Breuning 1967 from Şenköy Teşvikiye, Yalova, ABL= 21/25mm; ssp.

pseudomariettiSchweiger 1964 from Yeşigöller/Yığılca 1150m, Bolu, (leg. K.Staven), ABL = 17/22mm; ssp.

necopitanusBasquin & Darge 1986 from İsmet paşa, ABL= 17/19mm; D: ssp. charetianus Auvray 1992 from Demirköy, 395/640m, Est Edirne, (Col.D.Prunier), ABL= 19/23mm; spp. bischoffi

Chaudoir, 1848 from Yenigün, 800m, Samsun, (Col. B.Lassalle), ABL= 18-22mm; spp. ornatus Heinz, 1976 from Yaralıgöz (leg. Ph. Vingler), ABL= 17-20mm; spp. muchei Breuning, 1961 from Ilgaz Geçidi, Kastamonu, ABL = 19-21mm; spp. akensis

Haury, 1889 from Uluköy 1500m, Bursa, Uludağ, ABL= 20-23mm.

Geographical range: Turkey, northwestern Anatolia (Deuve 2004).

Chorotype: This distribution exemplifies a Northwestern Anatolian endemic chorotype.

Way of life and conservation: Species of wet forests and is not threatened by moderate logging.

Common. It requires the presence of a vegetative cover to withstand droughts.

Carabus Linné, 1758 (subg. Sphodristocarabus

Thomson, 1875)

Carabus (Sphodristocarabus) heinzi Breuning, 1964

Distribution s.lat. (all ssp. known): Turkey: Central and eastern Pontic Alps.

Specimens examined: Kumbet 1700m, Giresun, 1 specimen, ABL= 18-25 mm.

Geographical range: Turkey: central and eastern Pontic Alps. (Deuve 2004).

Chorotype: This distribution exemplifies a Northeastern Anatolian endemic chorotype.

Way of life and conservation: Species that lives mainly in forests and thickets, wet pastures reached altitude. Species not threatened by moderate logging.

Carabus (Sphodristocarabus) gilnickiiDeyrolle, 1869

Distribution s.lat. (all ssp. known): Turkey: Eastern Pontic Alps.

Specimens examined: Çamlık, 1800m, İkizdere, Rize, 1 specimen, ABL= 20-24mm.

Geographical range: Turkey: Eastern Pontic Alps (Deuve 2004).

Way of life and conservation: Species of wet forestsand and wet pastures, is not threatened by moderate logging.

Carabus (Sphodristocarabus) coruhneriensis

Cavazzuti, 1990

Distribution s.lat. (all ssp. known): Turkey: Eastern Pontic Alps.

Geographical range: Turkey: Eastern Pontic Alps (Deuve, 2004).

Way of life and conservation: Potential threats are overgrazing and forest degradation.

Carabus (Sphodristocarabus) enigmaticusHeinz, 1980

Distribution s.lat. (all ssp. known): Turkey: Eastern Anatolia.

Specimens examined: İspir, 1600m, Erzurum, 1 specimen, ABL= 27-29mm.

Geographical range: Turkey, eastern Anatolia (Deuve, 2004).

Carabus (Sphodristocarabus) georgia Cavazzuti, 1984

Distribution s.lat. (all ssp. known): Turkey, eastern central Anatolia: mountains south Erzincan (Deuve 2004).

Type locality: CE Turkey, about 10 km NE of Tunceli, 1100 m (Deuve 2004).

Geographical range: Turkey, eastern central Anatolia: mountains south Erzincan (Deuve 2004).

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Eastern Anatolian endemic chorotype.

Carabus (Sphodristocarabus) biroiCsiki, 1927

Distribution s.lat. (all ssp. known): Northern

Anatolia: near Samsun (Deuve 2004).

Type locality: N Turkey, Amasya (Deuve 2004). Specimens examined: Kavak Hacılar, 750m, Samsun, 1 specimen, ABL= 23-25 mm.

Geographical range: Northern Anatolia: near Samsun (Deuve 2004).

Way of life and conservation: Species of wet forests and is not threatened by moderate logging. It requires the presence of a vegetative cover to withstand droughts.

Common.

Carabus (Sphodristocarabus) pavesiiCavazzuti, 1992

Distribution s.lat. (all ssp. known): Turkey, northwestern Anatolia: Toprak (Vill. Izmir) (Deuve 2004).

Type locality: W Turkey, Izmir, Bornova. (Deuve 2004).

Geographical range: Turkey, northwestern Anatolia: Toprak (Vill. Izmir) (Deuve 2004).

Chorotype: This distribution exemplifies a Southwestern Anatolian endemic chorotype.

Carabus (Sphodristocarabus) macrogonus

Chaudoir, 1847

Distribution s.lat. (all ssp. known): Turkey, Northern Anatolia: Pontic Alps.

Specimens examined: spp. giresunensis Deuve 1992 from Zigana 1600m, Trabzon, ABL= 21/23mm, (leg. B.Lassalle); spp. schweigerinae

Schweiger, 1962 from Abant Gölü, ABL= 20-24mm (leg. Ph. Vingler); spp. resplendensBreuning, 1932 from Çayırkent 550m, Sarıbıyık, Samsun, ABL= 23-25mm; spp. drannadaghensis Breuning, 1964 from Drannaz Dag 1500m, Sinop, ABL= 21-23mm; spp. macrogonusChaudoir, 1847 from Zigana 1680m; ssp. pseudosimonianus Mandl 1975 from Gölköy 1900m, Ordu (leg. B.Lassalle), ABL= 21/25mm; ssp. keltepensis Taglianti & Bruschi 1976 from Keltepe 1500m, İzmit, (leg. B.Lassalle), ABL= 21/23mm; ssp. giachinoiCavazzuti 1991 from 15km Torull, 1500m, ABL= 21/23mm; ssp. simonianus

Breuning & Ruspoli 1970 from Mesudiye, (leg. B.Lassalle), ABL= 22/24mm; ssp. koserelensis Machard 1991 from Tirebolu 75m, Giresun, ABL= 23/25mm; ssp. lilianae Casale & Cavazzuti 1976 Fig. 9.Carabus (Sphodristocarabus) enigmaticusHeinz, 1980.

A: Habitus in dorsal view (A: from İspir 1600m, Erzurum, ABL = 27-29mm).

Fig. 10.Carabus (Sphodristocarabus) biroiCsiki, 1927. A: Habitus in dorsal view (A: from Kavak Hacılar 750m, Samsun, ABL = 23-25mm).

Fig. 7.Carabus (Sphodristocarabus) heinziBreuning, 1964. A: Habitus in dorsal view (A: from Kumbet 1700m, Giresun, ABL = 18-25mm).

Fig. 8.Carabus (Sphodristocarabus) gilnickiiDeyrolle, 1869. A: Habitus in dorsal view (A: from Çamlık 1800m, İkizdere, Rize, ABL = 20-24mm (leg. B.Lassalle)).

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from Golköy 1500m, Ordu, ABL= 22/24mm; ssp.

mollioniBreuning & Ruspoli 1976 from Zindan Dag 1300m, Sinop, (leg. B.Lassalle), ABL= 22/24mm; ssp. resplendens Breuning 1934 from Çayırkent 550m, Sarıbıyık, Samsun, ABL= 23/25mm; ssp.

akkusensisBreuning 1964 from (=ssp. guergentepeensis

Blumenthal 1975), Refahiye 1100m, Ordu (Col. D.Prunier), ABL= 20/25mm.

Geographical range: Northern Anatolia: Pontic Alps (Deuve 2004).

It requires the presence of a vegetative cover to withstand droughts. Common.

Carabus (Sphodristocarabus) giachinoi

Cavazzuti, 1991

Distribution s.lat. (all ssp. known): Turkey, northern Anatolia: Tersudagi Pass (vill. Gümüshane) (Deuve 2004).

Type locality: NE Turkey, Gümüshane, NE of Siran, Tersudagi Pass, 1800 m. (Deuve 2004).

Geographical range: Turkey, northern Anatolia: Tersudagi Pass (vill. Gümüshane) (Deuve 2004).

Chorotype: Unknown. Ecology: Unknown. Conservation: Unknown.

Carabus (Sphodristocarabus) tokatensis

Roeschke, 1898

Distribution s.lat. (all ssp. known): Turkey, Eastern central Anatolia.

Specimens examined: spp. simonae Cavazzuti, 1984 from Pülümür 1500m, Tunceli, ABL= 26-31mm; ssp. kosensisCavazzuti 1984 from Tersundağı Geç. 1925m, Gümüshane (Col. D.Prunier), ABL= 26/28 mm;spp. tokatensisRoeschke, 1898 from Yıldız Dağ 1400m, Tokat, (B.Lassalle), ABL= 19-23 mm. Geographical range: Turkey: Eastern central Anatolia (Deuve 2004).

Way of life and conservation: Species of concerned because of overgrazing in forests.

Conservation: Potential threats are overgrazing and forest degradation. It requires the presence of a vegetative cover to withstand droughts.

Fig. 11.Carabus (Sphodristocarabus) macrogonusChaudoir, 1847. A-N: Habitus in dorsal view (A: spp.

giresunensisDeuve 1992 from Zigana 1600m, Trabzon, ABL = 21/23mm, (leg. B.Lassalle); B: spp. schweigerinaeSchweiger, 1962 from Abant Gölü, ABL = 20-24mm (leg. Ph. Vingler); C: spp. resplendensBreuning, 1932 from Çayırkent 550m, Sarıbıyık, Samsun, 23-25mm; D: spp. drannadaghensisBreuning, 1964 from Drannaz Dag 1500m, Sinop, body long, 21-23mm; E: spp.

macrogonusChaudoir, 1847 from Zigana 1680m; F: ssp. pseudosimonianusMandl 1975 from Gölköy 1900m, Ordu (leg. B.Lassalle), ABL = 21/25mm; G: ssp. keltepensisVigna-Taglianti & Bruschi 1976 from Keltepe 1500m, Izmit, (leg. B.Lassalle), ABL = 21/23mm; H: ssp. giachinoiCavazzuti 1991 from 15km Torull, 1500m, ABL = 21/23 mm; I: ssp. simonianusBreuning & Ruspoli 1970 from Mesudiye, (leg. B.Lassalle), ABL = 22/24 mm; J: ssp. koserelensisMachard 1991 from Tirebolu 75 m, Giresun, ABL = 23/25mm; K: ssp. lilianae Casale & Cavazzuti 1976 from Gölköy 1500m, Ordu, ABL = 22/24mm; L: ssp. mollioni Breuning & Ruspoli 1976 from Zindan Dag 1300m, Sinop, (leg. B.Lassalle), ABL = 22/24mm, M: ssp. resplendensBreuning 1934 from Çayırkent 550m, Sarıbıyık, Samsun, ABL = 23/25mm; N: ssp. akkusensisBreuning 1964 from (=ssp. guergentepeensisBlumenthal 1975), Refahiye 1100m, Ordu (Col. D.Prunier), ABL = 20/25mm).

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Carabus (Sphodristocarabus) karasudominus

Cavazzuti, 1991

Distribution s.lat. (all ssp. known): Turkey, Eastern central Anatolia: Karasu Dagları (Deuve 2004).

Type locality: CE Turkey, Erzurum, Karasu Daglari 2100 m. (Deuve 2004).

Geographical range: Turkey, Eastern central Anatolia: Karasu Dagları (Deuve 2004).

Way of life and conservation: Potential threat is overgrazing.

Carabus (Sphodristocarabus) basquini Darge, 1990

Distribution s.lat.: East of Turkey: near Van Lake. s.lat.

Type locality: SE Turkey, Bitlis, between Resadiye and Gevas, Alacabük Dag. (Deuve 2004).

Geographical range: East of Turkey, near Van Lake. (Deuve 2004).

Chorotype: Unknown.

Carabus Linné, 1758 (subg. Tribax Fischer von Waldheim, 1817)

Carabus (Tribax) merdeniki Cavazzuti et Korell, 1992

Distribution s. lat.: Eastern Turkey: Göle (Deuve 2004).

Type locality: NE Turkey, Kars, near Göle. (Deuve 2004).

Geographical range: Eastern Turkey: Göle (Deuve 2004).

Common.

Carabus (Tribax) curlettiiCavazzuti, 1984

Distribution s. lat.: Eastern Anatolia, Sarıkamış. Type locality: CE Turkey, Kars, about 10 km SW Sarikamis 2000 m. (Deuve 2004).

Fig. 13.Carabus (Lamprostus) spinolaiDe Cristoforis & Jan, 1837. Habitus in dorsal view A-F: A: ssp. lohsei

Heinz & Blumenthal 1978 from Zindan Dag 1200m, Sinop (leg. B.Lassalle) (photo D.Prunier), ABL = 29/35mm; B: ssp. coeruleoviolaceusBreuning & Ruspoli 1970 from Boyabat 1300m, Sinop, (leg. B.Lassalle) (photo D.Prunier), ABL = 24/27mm; C: ssp. rouzieiBreuning & Ruspoli 1970 from Kurbagalı (photo D.Prunier), ABL = 28/33mm; D: ssp. usurpatorCavazzuti 1987 from Devrekani 1250m, Kastamonu, (leg. B.Lassalle) (photo D.Prunier), ABL = 28/32 mm; E: ssp. pseudescherichiBreuning 1962 from

Şereflikochisar, 1000m, Ankara (leg. B.Lassalle) (photo D.Prunier), ABL = 32/38mm; F: ssp. spinolaiCristoforii & Jan 1837 from Meçitozu, (photo D.Prunier), ABL = 32/37mm.

Fig. 12.Carabus (Sphodristocarabus) tokatensisRoeschke, 1898. A-C : Habitus in dorsal view (A: spp. simonae Cavazzuti, 1984 from Pülümür 1500m, Tunceli, ABL = 26-31mm; B: ssp. kosensis Cavazzuti 1984 from Tersundağı Geç. 1925m, Gümüshane (Col. D. Prunier), ABL = 26/28mm; C: spp. tokatensis Roeschke, 1898 from Yıldız Dağ 1400m, Tokat, (B.Lassalle), ABL = 19-23mm.

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Geographical range: Eastern Anatolia: Sarikamis (Deuve 2004).

Carabus Linné, 1758 (subg. Cechenochilus

Motschulsky, 1846)

Carabus (Cechenochilus) lazorum Belousov & Zamotailov, 1999

Distribution s.lat. : only one station, NE Turkey, S of Rize, Okçular Dagh, W of Demi Dagh, 2200 m (Deuve 2004).

Type locality: NE Turkey, S of Rize, Okçular Dagh, W of Demi Dagh, 2200 m (Deuve 2004).

Geographical range: Northeastern Turkey. Chorotype: This distribution exemplifies a Northeastern Anatolian endemic chorotype.

Way of life and conservation: High altitude species; does not seem in danger.

Carabus Linné, 1758 (subg. Lamprostus

Motshulsky, 1865)

Carabus (Lamprostus) spinolaiDe Cristoforis & Jan, 1837

Distribution s.lat. (all ssp. known): Northern and Southern Anantolia.

Type locality: N Turkey "Amasya". (Deuve 2004). Specimens examined: Çorum, TR, 1 specimen, 32-37 mm.

ssp. lohsei Heinz & Blumenthal 1978 from Zindan Dag 1200m, Sinop (leg. B.Lassalle) (photo D.Prunier), ABL= 29/35mm; ssp. coeruleoviolaceus

Breuning & Ruspoli 1970 from Boyabat 1300m, Sinop, (leg. B.Lassalle) (photo D.Prunier), ABL= 24/27mm; ssp. rouziei Breuning & Ruspoli 1970 from Kurbagalı (photo D.Prunier), ABL= 28/33mm; ssp. usurpator Cavazzuti 1987 from Devrekani 1250m, Kastamonu, (leg. B.Lassalle) (photo D.Prunier), ABL= 28/32 mm; ssp.

pseudescherichi Breuning 1962 from Sereflikochisar, 1000m, Ankara (leg. B.Lassalle) (photo D.Prunier), ABL= 32/38mm; ssp. spinolaiCristoforii & Jan 1837 from Mecitozu (photo D.Prunier), ABL = 32/37mm.

Geographical range: Turkey: Northern and Southwestern Anatolia (Deuve 2004).

Common.

Fig. 14.Carabus (Lamprostus) erenleriensisSchweiger, 1962: A-E: Habitus in dorsal view (A: spp. bithynicus Heinz et Korge, 1967 from Akcakoca, 600m, Bolu, (leg. B.Lassalle) (photo D.Prunier), ABL = 34/41mm; B: spp. ahmetustaensisCavazzuti, 1987 from Ahmetusta, 1000m, Zonguldak, (leg. B.Lassalle) (photo D.Prunier), ABL = 32/39mm; C: ssp. erenleriensisHeinz & Korge 1967 from Abant 1200m Bolu (leg. B.Lassalle) (photo D.Prunier),

ABL = 33/37mm; D: ssp. fortepunctatusBattoni & Blum 1973 from Mengen 700m, Bolu (leg. B.Lassalle) (photo D.Prunier), ABL = 30/35 mm; E: ssp. karadagensisBattoni & Blumenthal 1973 from Yurukyeri 700m, Karadag, Bolu (leg.

B.Lassalle) (photo D.Prunier), ABL = 33/40mm).

Fig. 15.Carabus (Lamprostus) robustusDeyrolle, 1869. A-D: Habitus in dorsal view (A: ssp. pseudocolossus Cavazzuti 1993 from Karabulduk, Giresun (leg.

B.Lassalle) (photo D.Prunier), ABL = 35/41mm; B: ssp. triumphansCavazzuti 1987 from Tirebolu 100m Giresun, (leg; B.Lassalle) (photo

D.Prunier), ABL = 40/45mm; C: ssp. singanensis Breuning, 1932 from Zigana 1680 m); D: ssp. robustusDeyrolle 1869 from Düzköy Yaylası, 1400/1700 m, 30 km Akçaabat, Trabzon (photo and Col. D.Prunier), ABL = 38/42mm and E: biotope (E: Zigana 1680m june 2006 biotope, (photo D.Prunier)).

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Carabus (Lamprostus) erenleriensisSchweiger, 1962

Distribution s.lat. (all ssp. known): Western Pontic Alps.

Type locality: NW Turkey, Bolu, Abant. (Deuve, 2004).

Specimens examined: spp. bithynicus Heinz et Korge, 1967 from Akcakoca, 600m, Bolu, (leg. B.Lassalle) (photo D.Prunier), ABL= 34/41mm; spp. ahmetustaensisCavazzuti, 1987 from Ahmetusta, 1000m, Zonguldak, (leg. B.Lassalle) (photo D.Prunier), ABL= 32/39mm; ssp. erenleriensis Heinz & Korge 1967 from Abant 1200m Bolu (leg. B.Lassalle) (photo D.Prunier), ABL= 33/37mm; ssp. fortepunctatus Battoni & Blum 1973 from Mengen 700m, Bolu (leg. B.Lassalle) (photo D.Prunier), ABL= 30/35mm; ssp. karadagensis

Battoni & Blumenthal 1973 from Yurukyeri 700m, Karadag, Bolu (leg. B.Lassalle) (photo D.Prunier), ABL= 33/40 mm.

Geographical range: Asiatic Turkey, Western Pontic Alps (Deuve 2004).

Chorotype: This distribution exemplifies a Northwestern Anatolian endemic chorotype.

Common.

Carabus (Lamprostus) robustusDeyrolle, 1869

Distribution s.lat. (all ssp. known): Pontic Alps, east of Giresun.

Type locality: N Turkey, near Trabzon, Elleon (Deuve 2004).

Specimens examined: Zigana, 1680 m TR, 1 specimen.

Geographical range: Asiatic Turkey: Pontic Alps, east of Giresun (Deuve 2004).

Chorotype: This distribution exemplifies a North Turkey endemic chorotype.

This species also inhabits culture hazel. It is vulnerable to pesticide application.

Carabus (Lamprostus) chalconatus

Mannerheim, 1830

Distribution s.lat. (all ssp. known): Turkey, eastern and central Anatolia.

Type locality: NE Turkey, Bayburt, Kop Dag. (Deuve 2004).

Specimens examined: sp. chalconatus

Mannerheim, 1830 from İspir 1600m, Erzurum,

(photo D.Prunier), ABL= 26-27mm.

Geographical range: Turkey, central and eastern Anatolia (Deuve 2004).

Way of life and conservation: Forests, and lowland and upland pasture, and wastelands. It is not threatened in natural habitats. A common species, but absent from areas of intensive farming.

Fig. 17.Carabus (Procrustes) paiafaWhite, 1845: A-B: Habitus in dorsal view (A: spp. paiafaWhite, 1845 from Baba Dağ 1000-1300m, Fethiye, ABL = 37-48mm; B: spp. acuticollisMotschulsky, 1858 from Karapınar 1248m, Alanya, (Col. D.Prunier), ABL = 35-40 mm; C, D, E: biotope(C: from Fethiye Babadag payafa, Mugla; D: from Ölüdeniz payafa,

Mugla; E: from Irmasan 1200m, Alanya). Fig. 16.Carabus (Lamprostus) chalconatusMannerheim,

1830. A: Habitus in dorsal view (A: sp. chalconatus Mannerheim, 1830 from İspir 1600m, Erzurum, (photo D. Prunier), ABL = 26-27mm).

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Species of concern from overgrazing in forests and pastures.

Carabus Linné, 1758 (subg. ProcrustesBonelli, 1809)

Carabus (Procrustes) paiafaWhite, 1845

Distribution s.lat. (all ssp. known): southern Anatolia, Taurus, Lycia.

Type locality: S Turkey, "Lycia, Günek". (Deuve 2004).

Specimens examined: spp. paiafa White, 1845 from Baba Dag 1000-1300m, Fethiye, ABL= 37-48 mm; spp. acuticollis Motschulsky, 1858 from Karapınar 1248m, Alanya, (Col. D.Prunier), ABL= 35-40 mm.

Geographical range: Turkey, southern Anatolia: Taurus, Lycia (Deuve 2004).

Way of life and conservation: Species of concern from overgrazing in forests. It requires the presence of a vegetative cover to withstand droughts. Species hélicophage very sensitive to the presence of Snail. Requires a rocky habitat.

Carabus (Procrustes) anatolicusChaudoir, 1857

Distribution s.lat. (all ssp. known): Cyprus and southern Anatolia. (Deuve 2004).

Type locality: Cyprus. (Deuve 2004).

Specimens examined: spp. anatolicus Chaudoir 1857 from Trodos 1600m Chyprus, ABL= 26/29mm; ssp. lycicus Breuning 1934 from Alanya, (Col. D.Prunier), ABL= 28/31mm.

Geographical range: Cyprus and southern Anatolia (Deuve 2004).

Chorotype: This distribution exemplifies a Southwestern Anatolian endemic chorotype.

Way of life and conservation: Forests, and lowland and upland pasture, and wastelands. It is

not threatened in natural habitats. A common species, but absent from areas of intensive farming. Species of concern from overgrazing in forests.

Carabus (Procrustes) mulsantianus Morawitz, 1886

Distribution s.lat. (all ssp. known): Southeastern Anatolia and Syrian border (Deuve 2004).

Type locality: SE Turkey, near Adana. (Deuve 2004).

Specimens examined: ssp. aleppensis Lapouge 1914 from Akpınar N. Reyhanil, Hatay, (Col. D. Maquet) (photo Noël Mal), ABL= 30/34mm;ssp.

akbesianus Lapouge 1914 from Akyokus 1100m, Gaziantep, (leg. B.Lassalle), ABL= 28/32 mm; ssp.

bernhauerorum Heinz 1977 from Kücüksu 1400m, Göksun Kahramanmaraş, (Col.D.Prunier), ABL= 34/38mm; ssp. mulsantianus Morawitz 1886 from Kahramanmaraş 790m, (Col. D.Prunier), ABL= 29/35mm; ssp. nurdagensis Blumenthal 1967 from Atik Aylası 850/1100m Belen, Nurdag, Hatay, (Col. D.Prunier), ABL= 32/35mm.

Geographical range: Southeastern Anatolia and Syrian border (Deuve 2004).

Way of life and conservation: Forests, and lowland and upland pasture, and wastelands. It is not threatened in natural habitats. A common species, but absent from areas of intensive farming. Species of concern from overgrazing in forests and Fig. 18.Carabus (Procrustes) anatolicusChaudoir, 1857: A, B:

Habitus in dorsal view (A: spp. anatolicusChaudoir 1857 from Trodos 1600m Chyprus, ABL = 26/29mm; B: ssp. ssp. lycicus Breuning 1934 from Alanya, (Col. D.Prunier), ABL = 28/31mm; C:

biotope (from Irmasan 1200m, Alanya). Fig. 19.Carabus(Procrustes) mulsantianusMorawitz, 1886.

A-D: Habitus in dorsal view (A: ssp. aleppensis Lapouge 1914 from Akpınar, Reyhanlı, Hatay, (Col.D.Maquet)(photo Noël Mal), ABL = 30/34mm; B: ssp. akbesianusLapouge 1914 from Akyokuş 1100m, Gaziantep, (leg. B.Lassalle), ABL = 28/32 mm; C: ssp. bernhauerorumHeinz 1977

from Kücüksu 1400m, Göksun Kahramanmaraş, ABL = 34/38mm; D: ssp. mulsantianusMorawitz 1886 from Kahramanmaraş 790m, ABL = 29/35mm; E: ssp. nurdagensisBlumenthal 1967 from Atik Aylası 850/1100m Belen, Nurdag, Hatay,

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pastures.

Carabus Linné, 1758 (subg. ApoplesiusDeuve, 1990)

The observation of this sub-genre is difficult because the insects come out in conditions of humidity and temperature. They only run a few days a year on surface and remain buried for the rest of time in the crannies of the soil.

Carabus (Apoplesius) deuveiLassalle, 1986

Distribution s. lat.: East of Turkey : between Erzincan and Tunceli (Deuve 2004).

Type locality: SE Turkey, Tunceli, Pülümür (Deuve 2004).

Geographical range: East of Turkey: between Erzincan and Tunceli (Deuve 2004).

Way of life and conservation: The species inhabits protected rocky biotopes, and is rare, but not threatened.

Carabus (Apoplesius) pseudodepressusMachard, 1988

Distribution s. lat.: East of Turkey: southern bank of Van Lake (Deuve 2004).

Type locality: SE Turkey, Bitlis, Resadiye (Deuve, 2004).

Geographical range: East of Turkey: southern bank of Van Lake (Deuve 2004).

Carabus (Apoplesius) legrandiDeuve, 1991

Distribution s. lat.: Eastern Anatolia: near Kars (Deuve 2004).

Type locality: E Turkey, Kars, Digor. (Deuve, 2004).

Geographical range: Eastern Anatolia: near Kars (Deuve 2004).

Way of life and conservation: The species inhabits protected rocky biotopes, and is rare, but

not threatened.

Carabus (Apoplesius) omphreodesReitter, 1898

Distribution s. lat.: East of Turkey: near Mardin (Deuve 2004).

Type locality: SE Turkey, Mardin (Deuve, 2004). Specimens examined: In private collection, not available for the study.

Geographical range: East of Turkey: near Mardin (Deuve 2004).

Carabus Linné, 1758 (subg. MegodontusSolier, 1848)

Fig. 20.Carabus(Megodontus) bonvouloiriChaudoir, 1863. A-E: Habitus in dorsal view (A: spp. fortunantus Heinz, 1970 from Hopa 800m, Artvin, ABL = 36-44 mm; B: spp. bonvouloiriChaudoir, 1863 from Tamdere, 1400-1550m, Giresun, ABL = 30-40mm; C: ssp. kaschkarensisBlumenthal & Breunig 1967 from Ayder 1600m, Rize, ABL = 30/39 mm (leg. B.Lassalle); D: ssp. casaleiCavazzuti 1986

from Kabaca, ABL = 32/37mm; E: ssp.

holatdaghensisBreuning 1967 from Uluköy 1500m, Torul, Gümüshane, (leg. B.Lassalle), ABL = 34/40

mm; and F-H: biotope (F: from Hopa, Yolder Basob 85m, biotope Lipaster; G: from Zigana 1900m, june 2006; H: from Col de Zigana 1900m,

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Carabus (Megodontus) bonvouloiri Chaudoir, 1863

Distribution s. lat.: Turkey, northern Anatolia (Deuve 2004)

Type locality: NE Turkey near Trabzon (Deuve, 2004).

Specimens examined: spp. fortunantus Heinz, 1970 from Hopa 800m, Artvin, ABL= 36-44 mm; spp. bonvouloiri Chaudoir, 1863 from Tamdere, 1400-1550m, Giresun, ABL= 30-40mm; ssp.

kaschkarensis Blumenthal & Breunig 1967 from Ayder 1600m Rize, ABL= 30/39 mm (leg. B.Lassalle); ssp. casaleiCavazzuti 1986 from Kabaca, ABL= 32/37mm; ssp. holatdaghensisBreuning 1967 from Uluköy 1500m, Torul, Gümüshane, (leg. B.Lassalle), ABL= 34/40 mm.

Geographical range: Turkey, northern Anatolia (Deuve 2004).

Common.

DISCUSSION

Endemic species are of both local and global importance. They increase the amount of genetic diversity and offer insight into biogeographical questions, such as where certain species originated and how distribution patterns have changed over time (Skarberk 2008). There are some researches on the listing, ecology, and conseraviton of endemic carabids conducted in different countries (Gueorguiev 2007, Abdel-Dayem 2012, Boieiro et al. 2012). Researhes about endemic species of

Carabus as indicator and biological control agents are very important for the next studies (Boieiro et al. 2012).

Many endemic species occupy very limited habitats and have small populations but have not been listed as threatened if their environments or populations have not declined. But endemic species have been affected negatively for different reasons like the adverse effects of human activity and climatic factors nowadays, and some of them may be under theat.

One of most important studies on the threatened Coleoptera is is called “European Red List of Saproxylic Beetles” (Nieto and Alexander 2010, Anonymous 2012). The European Red List of Saproxylic Beetles was published by the

International Union for Conservation of Nature (Anonymous 2012). A lot of Saproxylic Beetle species are classified in different threatened categories in the European Red List. (Nieto and Alexander 2010, Anonymous 2012). A similar list was prepared for the Turkish species (Avgın et al. 2014). The list of Turkish Saproxylic Beetles threatened was prepared using the paper on the European Red List of Saproxylic Beetles (Avgın et al. 2014). But these lists do not include carabids.

Moreover, there are some papers on the threatened carabids and conservation of them in Europe. In some of these, researches recorded that many species of carabids are threatened by habitat fragmentation (Niemela, 2001). For instance, Desender & Turin (1989) list 142 carabid species that are endangered in western Europe (Denmark, Netherlands, Belgium, and Luxembourg) represen-ting as much as one third of the carabid fauna in these countries. At least some of them are threatened because of habitat fragmentation.

But research has not been preformed on the endemic carabids and threatened species of carabids in Turkey so far. This paper is the first comprehensive study on the endemic carabinae in Turkey. The information obtained from this paper can be use in ecological, biological control, agriculture, and forest studies. Moreover, this paper provides a contribution to the prepare endemic species list, threated species list, and to the preservation of biodiversity.

The General State of Turkish Carabinae is Presented Below Based on our Previous Observations and Comments:

In the foregoing annotations of endemic species of the Carabus spp. in Turkey, personal field observations and other information from the literature on Turkey’s endemic species of Carabus

lead to the conclusion that they can serve as indicator species of biotope health. The history of these beetles includes ancient Turkey when forests were larger and denser than today and a low human pressure resulted in a set of stable natural environments required by the species of Carabus. Such stability in many regions has deteriorated considerably as human colonization greatly changed the appearance of landscapes in Turkey due to agriculture and forestry practices.

While in much of the western part of Europe many landscapes have been irreversibly changed,

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Turkey has a chance to avoid that unwelcomed possibility in many parts of this large country. Turkey has a large variety of climates including a continental climate and a mild climate along the Mediterranean coast. There are also very high mountains with alpine climate, plateaus with dry, or desert climates, and in each of these are species of

Carabus adapted to the biotopes.

Unfortunately, in several areas of Turkey overgrazing in both forests and pastures has changed the nature of soils (for example, in the south at Lycian Taurus/Kahramanmaras). Domestic ungulates even eat seedling trees, so regeneration of forests is compromised. In the Baba Mountain (Fethiye, Lycian) and Irmasan Gecidi (Antalya) there are beautiful forests whose future is jeopardized by the repeated passage of domestic herds that leave trampled bare soil.

In northern Turkey, heavy rains allow rapid recovery of vegetation cover and beetles are still abundant. However, in the region of Trabzon and Hopa, the lower altitude forests are encroached upon by intensive cultivation of tea plantations and natural forests there are being completely replaced. On the coast of Trabzon, forest undergrowth of rhododendrons is being replaced by the intensive cultivation of hazelnuts (Corylus). And, in general, throughout the agricultural lands chemical pesticides are used heavily and this adds to the human pressure on the species of Carabus and all other wildlife, as well as the people who apply the chemical.

Adults of Carabus (subgenus Procerus) have two particular sensitivities. They are very large, diurnal, and very visible. And, they are often victims of crushing by the public. They are avid consumers of land snails (Helix lucorum L.) and require forest cover. Overgrazing, which removes the Kermes Oak (Quercus cocciferaL.) deprives them of the shelter that allows survival during heat waves in the south of Turkey (in the vicinity of Harbiye, and the Amanos

and Taurus mountains).

While the biological potential in Turkey is still

enormous with exceptional species richness, only sound landscape management will keep it that way and prevent species from becoming threatened, or driven to extinction. It is therefore urgent to establish and maintain in perpetuity a network of preserved areas that can buffer against the loss of wildlife and the endemic species of Carabus

documented above.

It is important to keep track of these species. In conducting regular crops in the field, we can evaluate the maintenance or decline of wildlife. We must keep a photographic record of the media surveyed in order to establish a correlation between the state of the environment and the presence of the species studied. It would be very interesting to involve professionals or amateur entomologists to the observation of species. Sharing their results, they can contribute to the monitoring of species. We must therefore ask the local authorities to help entomologists in the field. It should also take into account the climatic factor and rainfall, which can greatly affect the results. The Carabus species are subject to natural and artificial conditions combi-ned. Human impacts may worsen temporarily the difficult natural conditions (drought and heat) that would have been tolerated without alteration of the natural environment. In the strictly forested areas, clear-cutting would be prohibited. Because, it causes large temperature differences typically unbearable for species in the undergrowth, drying of the humus, and erosion with degradation of surface horizons. This leads towards destruction of the microfauna of the soil surface. Another scourge is often noticed in the territory that is the deposits of detritus in the natural environment: cans, cups, plastic bottles, bags, and a variety of other garbage. The practice of throwing waste into the environment contributes to impoverishing nature.

ACKNOWLEDGEMENTS

We heartily thank Dr. Terry L. Erwin (Smithsonian Institution, Washington, DC) who greatly improved the original version of this manuscript.

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