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The diversity of cave crickets (Orthoptera, Rhaphidophoridae) remarkably differs between geographic regions in Anatolia. While only 4 species are distributed in northern Anatolia where Black Sea climate prevails, 14 species are found throughout southern and western Anatolia where the Mediterranean climate is dominant. However, no cave crickets were reported from the middle and eastern Anatolia where the continental climate is present. There is no data-based study on the distribution pattern of cave cricket species in Anatolia. This study aimed to reveal any possible relation between climatic conditions and distribution of cave crickets in three caves selected as representatives for three climate types present in Anatolia. Temperature and relative humidity in the ecological zones of the given caves and the surface were periodically measured by using data loggers. Our data show within cave variations in temperature and relative humidity among all three caves: (i) variations in temperature and relative humidity decreased from the entrance zone to the dark zone, (ii) over the winter period, temperature increased from the entrance zone to the dark zone, (iii) over the summer period, temperature decreased from the entrance zone to the dark zone, (iv) the most distinct temperature variation was measured in continental climate, and (v) the major similarity of climatic conditions between cave and climate type was observed in the Black Sea region which cave crickets species have widened distribution and limited speciation.
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A Comparative Study on Temperature and Relative Humidity

                                                         

Mehmet Sait TAYLAN1 Nadim YILMAZER2 Deniz 3*
1Department of Biology, Institute of Science, Hakkari University, Hakkari, TURKEY,
2,3Department of Biology, Faculty of Art and Science, Namk Kemal University, Tekirda, TURKEY
e-mails: msaittaylan@hakkari.edu.tr, nyilmazer@nku.edu.tr, denizsirin19@gmail.com
ORCID IDs: 10000-0002-2084-920X, 20000-0002-9935-9608, 30000-0001-5475-173X
ABSTRACT

            
            

the continental climate is present. There is no data-based study on the distribution pattern of cave cricket
       
distribution of cave crickets in three caves selected as representatives for three climate types present in
Anatolia. Temperature and relative humidity in the ecological zones of the given caves and the surface

and relative humidity among all three caves: (i) variations in temperature and relative humidity decreased
                
entrance zone to the dark zone, (iii) over the summer period, temperature decreased from the entrance

        

Key words: Rhaphidophoridae, cave crickets, cave zones, climatic factors, distribution, Anatolia.
J. Entomol. Res. Soc., 22(1): 53-73, 2020 Research Article
Print ISSN:1302-0250 Online ISSN:2651-3579
(2020). A Comparative study on temperature and relative humidity

crickets (Insecta, Orthoptera, Rhaphidophoridae). Journal of the Entomological Research Society,
22(1), 53-73.
54
TAYLAN, M.S., YILMAZER, N., ŞİRİN, D.

Caves, naturally occurring underground spaces, range in size from single small
           
host many unique biota that are highly interconnected and interdependent, and that

      

of as constant, they directly depend upon the surrounding surface environment that is

such as temperature, relative humidity, and light intensity, these conditions can change



Caves are valued as natural underground laboratories by scientists since they
can provide insight into evolutionary and ecological processes because of their
        


 



advances in speleology have enabled the development of biospeleology involving
2


 






          
(reproduction, nutrition, physiology, etc.) of cave crickets have been restricted to

crickets in Anatolia. The distribution of cave crickets (Orthoptera, Rhaphidophoridae)


irin, 2015), 14 species
55
A Comparative Study on Temperature and Relative Humidity Data of Cave Crickets



       

2, Fig. 1). Although some previous studies assumed that ancient central Anatolian lake





model organisms in three caves selected to represent three climate types seen in
Anatolia (Fig. 1), (i) by comparing the annual data of temperature and RH for each
  

the biogeographic distribution of cave crickets.
Table 1. The data of surface and ecological zones of the study caves.
Caves Measured data Surface Entrance
zone

zone
Dark
zone

Annual mean temperature 18.98 15.06 15.65 16.64
 50.64 98.53 98.91 98.94
 32.99 20.83 17.2 17.61
 5.06 7.78 14.22 15.06
 77.8 99.0 99.0 99.0
 18.37 97.0 97.0 98.0

Annual mean temperature 11.10 9.79 10.08 9.76
 78.65 78.61 99.85 98.98
 23.6 15.14 12.1 10.72
 -0.98 6.25 8.74 8.89
 91.75 88.61 99.99 99.0
 60.79 68.71 96.61 97.8

Annual mean temperature 11.81 11.02 9.43 10.09
 57.12 98.60 98.92 98.99
 30.23 16.98 10.6 10.56
 -7.83 5.1 7.35 9.56
 87.25 99.0 99.0 99.0
 27.29 97.0 97.08 98.99
56
TAYLAN, M.S., YILMAZER, N., ŞİRİN, D.
Fig. 1. Localities of the study caves and climatic regions in Turkey. The spots indicate the locations of the
cave crickets.



The climate mainly determined by the topography and other geographical factors





are especially severe. Mediterranean climate prevails in the coastal areas of Turkey

  





mm for the central Anatolia, 321 and 1230 mm for the East Anatolia, and 331 and


annual mean temperature (AMT) in Turkey varies from 3.6°C to 20.1°C, reaching a


is around 648 mm, ranging from 295 (the central Anatolia) to 2220 mm (the eastern


57
A Comparative Study on Temperature and Relative Humidity Data of Cave Crickets


Mediterranean, the Black Sea and the Continental climate regions, respectively (Fig. 1).



        


      

Dolichopoda lycia

Sipahiler cave having dimensions of 338 m length and 32 m depth is situated (N


investigation of the cave has not yet been carried out, the cave is famous for its
    

cave in 2009. Troglophilus aspegi
(Taylan et al, 2015).




and have a thickness of 30-40 m. This cave has almost completed its development
process, 



     
to continuously monitor air temperature and relative humidity (RH) at one-hour
         
placed in environments such as caves and cold storages/rooms. They measure the







58
TAYLAN, M.S., YILMAZER, N., ŞİRİN, D.





        

    
    



of temperature and humidity values of each cave. Besides, descriptive statistical



          
  


study cave.


the climate types as each cave is in a separate climate) compared to climatic
     



   


        



   
 



59
A Comparative Study on Temperature and Relative Humidity Data of Cave Crickets


in the remainder of the year (Table 1, Fig. 2A).

characteristics.
Sum of Squares df Mean Square F p
Surface
 1972,862 2 986,431 12,980 ,000
 11627,119 153 75,994
Total 13599,980 155
Entrance zone
 793,314 2 396,657 28,810 ,000
 2106,494 153 13,768
Total 2899,808 155

 1218,735 2 609,367 686,338 ,000
 135,841 153 ,888
Total 1354,576 155
Dark zone
 1564,857 2 782,428 2066,285 ,000
 57,936 153 ,379
Total 1622,792 155

   

°C in 
cave, 11.02 °C in °C in Sipahiler cave (Table 1). Although
      
entrance zone of all three study caves. Minimum values of 7.78 °C, 5.1 °C, and 6.25°C

and Sipahiler caves, respectively (Table 1, Fig. 2B). The WMTs gradually increased






°C in °C in Sipahiler

higher than those of the entr




60
TAYLAN, M.S., YILMAZER, N., ŞİRİN, D.


of all three caves from 2.98 °C to 3.36 °C (Fig. 2C).


Dependent





Mean


pDependent





Mean


p
Surface 
1,00
2,00 ,000 ,000

zone 
1,00
2,00 ,000 ,000
3,00 ,000 ,000 3,00 ,000 ,000
2,00
1,00 ,000 ,000
2,00
1,00 ,000 ,000
3,00 ,917 ,917 3,00 ,003 ,003
3,00
1,00 ,000 ,000
3,00
1,00 ,000 ,000
2,00 ,917 ,917 2,00 ,003 ,003
Entrance
zone 
1,00
2,00 ,000 ,000
Dark
zone 
1,00
2,00 ,000 ,000
3,00 ,000 ,000 3,00 ,000 ,000
2,00
1,00 ,000 ,000
1,00
1,00 ,000 ,000
3,00 ,245 ,245 3,00 ,025 ,025
3,00
1,00 ,000 ,000
3,00
1,00 ,000 ,000
2,00 ,245 ,245 2,00 ,025 ,025



61
A Comparative Study on Temperature and Relative Humidity Data of Cave Crickets


 






       
, and
gradually increased until the end of the year (Fig. 2D). In this zone, the minimum WMTs







        
         

      






 





    


(a drop from 17.61 °C to 17.25 °C), and subsequently another temperature drop of



gradual increase until the end of the year (Fig. 3A).
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TAYLAN, M.S., YILMAZER, N., ŞİRİN, D.

    

   
          












   








          
          
,
    


63
A Comparative Study on Temperature and Relative Humidity Data of Cave Crickets
ecological zones (Table 7). The WMTs of the atmospheric and entrance zones of this




     
dropped during the remainder of the year, reaching the initial values (Fig. 3C). In the

  


in the WMTs, reaching a minimum value of 9.56°C. The WMTs gradually increased
during the remainder of the year (Fig. 3C).


among the caves (or among the climate types as each cave is in a separate climate)
         




climate characteristics.
Sum of
Squares df Mean Square F p
Surface
 22367,091 2 11183,546 57,719 ,000
 29644,926 153 193,758
Total 52012,018 155
Entrance
zone
 13807,776 2 6903,888 925,015 ,000
 1141,922 153 7,464
Total 14949,698 155

 30,066 2 15,033 85,221 ,000
 26,989 153 ,176
Total 57,055 155
Dark zone
 ,087 2 ,044 1,573 ,211
 4,239 153 ,028
Total 4,326 155
         


     
64
TAYLAN, M.S., YILMAZER, N., ŞİRİN, D.
  















   
  
65
A Comparative Study on Temperature and Relative Humidity Data of Cave Crickets
RH of this zone remained constant around 99.99% in Sipahiler cave throughout the
       nd 


terms of dark zone RH values (Table 6). A constant RH of 99% prevailed in this zone
D).

         







nearly constant at a value of 97-99% throughout the year (Fig. 5A).


    




dark zones, respectively (Fig. 5B).
   





  
remained constant at a value of 99% in the dark zone (Fig. 5C).

all three caves: (i) variations in temperature and RH decreased from the entrance zone

the dark zone, (iii) over summer period, temperature decreased from the entrance to




66
TAYLAN, M.S., YILMAZER, N., ŞİRİN, D.

s Cave).



           
temperature and RH data of the zones of the given caves are generally similar in the





A study by Bekaro olu (2010) provided annual temperature values




1550 m is geographically in the Black Sea Region, its location is closer to the Eastern



      




zone 10.09 °C) (Bekaroolu, 2010). Once again concerning Karaca
 

o

67
A Comparative Study on Temperature and Relative Humidity Data of Cave Crickets

lighting, electricity cabling, tourist paths, and open to visitors for 8 months of the year.
When the number of species and geographical distribution of cave crickets in
Turkey is taken into account, a remarkable distribution is noted: There are four species
(Dolichopoda euxina Semenov 1901, Dolichopoda noctivaga
2007, Troglophilus tatyanaeTroglophilus aspegi 

Dolichopoda aranea Bolivar 1899, D. pusilla Bolivar 1899, D.
sbordoniiD. lyciaD. sutini 
Taylan 2012, D. fortuitaTroglophilus escalerai Bolivar, 1899,
T. adamovici Us 1974, T. gajaci Us 1974, T. bicakciiT.
alanyaensis     T. fethiyensis Taylan, Di
 T. ferzenensis 
2012, T. ozeli


   
irin, 2016) (Table 2).

Continental climate is dominant. Diversity and distribution patterns of species groups




nal, 2016) in Anatolian Orthoptera
have been proposed regardless of annual climatic data until this study.


     

zone of Sipahiler cave in the Black Sea climate region has an RH of 60-90% (Fig.





end of the year. Hence, if RH values of the three cave zones supported cave cricket

climate region. In this case, one can say that surface-climatic conditions (annual
         
distribution of cave crickets (Table 2).
68
TAYLAN, M.S., YILMAZER, N., ŞİRİN, D.
Table 2. Cave crickets, their localities and climatic zones in Anatolia.
 

 Black
Sea 
  + - -
 Hatay, Akbez Cave + - -
Semenov, 1901  - + -
 
cave + - -
 
cave + - -
 
 - + -
 
Cave + - -
, 2015  + - -
  + - -
Us, 1974  + - -
Us, 1974  + - -
 
 + - -
T. tatyanae   - + -


 + - -
      

 + - -


 + - -
 

 + - -
  - + -
*Data from Bolivar 1899, Semenov 1901, Us 1974,Rampini and Di Russo 2003, Di Russo et al. 2007,



droppings) as source of energy and nutrients is not found or occurs in small amounts

Therefore, more realistic scenarios about the diversity and distribution of cave crickets
can be produced by considering both in-cave conditions and surface climatic conditions

species (Dolichopoda lyciaTroglophilus aspegi
 cave does not contain any cave crickets species.
The most species density of D. lycia 
On the other hand, most species density of T. aspegi observed in the entrance zone
      
T. aspegi 
distribution zone than D. lycia species  
caves zones. The specimens of T. aspegi 
all zones and main galleriesD. lycia 
2006) have limited distribution in the side galleries
69
A Comparative Study on Temperature and Relative Humidity Data of Cave Crickets
From the data of this study, RH of the atmospheric zone of Sipahiler cave in the
Black Sea climate region appears to be very close to that surface of the cave, especially


region in Turkey (Doanay, 2004), the Black Sea Region generally has
a forest structure consisting of non-evergreen trees (Fagus sp., Carpinus sp., Alnus
spp. etc) at altitudes of 200 m to 1200 m, and coniferous trees resistant to harsh


     
Dolichopoda noctivaga


       
temperatures outside the cave (Fig. 2A, Fig. 3A) may dissuade cave crickets from their



arid-climate-loving Calabrian pine (Pinus brutia) forests at altitudes of 250-300 m to
Cedrus libani



   
among the populations, and correspondingly speciation.

some assumptions from this study based on data logging. Our study reveals that
distinct variations in temperature and RH lead to minute changes in the dark zone in
that a 50 ºC variation drops the temperature of the dark zone by 1-2 ºC, suggesting
that the cave ecosystem tolerates considerable changes in the outdoor atmosphere.




      
revealed by this study that temperature and RH gradually drop from the entrance

ecological zone (Fig. 3D, Fig. 5D). Concordantly, and the high tolerance of dark zone
to temperature variations makes caves important shelters for many invertebrates,
           

70
TAYLAN, M.S., YILMAZER, N., ŞİRİN, D.

      from The Society of Anatolian
 lu from Akdeniz University
urlu from Hacettepe University Cave



1929B011100092].
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
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European Journal of Entomology, 107, 631-645.
  Theoretical and
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         
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          
and vegetation from a snapshot in Mediterranean Anatolia: role of latitudinal position in altitudinal
Journal of Natural History, 44, 1343-1363.
     , B. (2010). Chorthippus brunneus subgroup (Orthoptera,

the lineage. Zootaxa, 2410, 1-28.
, B. (2011). Myrmeleotettix
     
end of the Anatolian refugium. Zootaxa, 2917, 29-47.
73
A Comparative Study on Temperature and Relative Humidity Data of Cave Crickets
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Akdeniz University, Antalya, Turkey.

 Turkey. Zootaxa, 3597, 33-40.

 -
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Troglophilus
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Troglophilus Krauss,
1879 (Orthoptera: Rhaphidophoridae) from northern Anatolia, Turkey. Journal of the Entomological
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Dolichopoda
of ancient central lake system. Insect Systematics and Evolution. 473, 267-283.
Ceuthophilus secretus
          American
Midland Naturalist, 154, 97-114.
            
Entomologist’s Monthly Magazine, 110, 182-193.
          
 Zootaxa, 4206, 1-23.
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Biospeologie: la biologie des animaux cavernicoles
            
Journal of Cave and Karst Studies, 74, 1-6.

of the cave cricket, Hadenoecus cumberlandicus. Journal of Cave and Karst Studies, 64(2), 140-144.
Received: April 24, 2019 Accepted: February 13, 2020
... The region Tuluntaş Cave is located in, is described with continental climate regime with sharply contrasting seasons in temperature. The research assessed the cricket populations of these caves whether a correlation exists with these conditions while the other parameters are also taken into consideration such as vegetation cover and elevation (Taylan, Yılmazer, & Şirin, 2020). Aside from locusts, arachnoids (Kunt, Özkütük, Elverici, Marusik, & Karakaş, 2016;Ribera, Elverici, Kunt, & Özkütük, 2014) millipede (Antić, Çetin, Turantepe, & Gürbüz, 2016;Enghoff, 2006), amphipoda (Özbek, 2007, 2012a, 2012bÖzbek & Oktar Guloglu, 2005;Özbek, Yurga, & Külköylüoǧlu, 2013;Özkan, 2009), fishes (Özdemir & Erkakan, 2014), ostracoda and planktons (Külköylüoğlu, Yavuzatmaca, Karacaoğlu, & Telli, 2014) were examined in various studies. ...
Chapter
Karstic lands cover the %40 of the total area of Turkey. Examination of these areas situated in the Alpine–Himalayan Orogenic Belt plays a key role in shedding a light on the tectonic evolution of this huge area that elongates from Europe to Asia, besides the tectonic evolution of Turkey. The research of the karstic lands exerts a great importance for the urban policies within the cities and villages settled in these geological regions, in order to require rigorous approaches to the engineering problems during the stages such as road and freshwater system construction or wastewater treatment. The most well-known karstic formations and caves exert a great impact on the development of landscapes as underground drains, since caves have been investigated in the context of geomorphological setting, lithology and geologic structure, groundwater chemistry and hydrology, aquifer type as well as regional tectonics, climate research, biology, microbiology, and in-cave environmental conditions. The historical background of the particular research for each selected caves from Turkey is given in Chap. 4 of this book. As comprehensively treated in Chap. 2, the state of the art in cave archaeology was excluded from the contents of this chapter.
Chapter
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Two life-cycle patterns can be identified in heterospecific or conspecific populations of Dolichopoda cave crickets. One pattern, typical of populations living in artificial caves, is characterized by strong seasonality in the age structure, egg and nymphal diapause and reproductive period. The second pattern, typical of populations inhabiting natural caves, is characterized by absence of such seasonality. These seasonal and aseasonal life cycles also involve demographic differences. In particular aseasonal populations have smaller clutches and larger eggs than seasonal populations. Moreover, studies on several populations revealed that the natural caves tend to hold smaller populations. Variation in the life-cycle traits of Dolichopoda populations does not fit any geographical cline, but appears to be related to ecological and historical features of individual caves. Particularly, the availability of guano, as a cave-restricted food resource, represents the main factor determining life-cycle variation. Such variation can be interpreted as adaptation to a particular coarse-grained discontinuous landscape, where different cave habitats favour the expression of different life-cycle types.
Thesis
In this research, taxonomy, phylogeny and biogeography of cave crickets which were collected between years 2006-2010 from different regions of Turkey were investigated in the light of morphological and molecular data. In this thesis which is the first study on the cave crickets of Turkey, morphologically, 19 quantitative characters of Troglophilus and 20 quantitative characters of Dolichopoda were measured and the taxonomy of cave crickets found in Turkey were also studied by examining qualitative characters used for their identification. Besides qualitative and quantitative characters of specimens in Caucasia and Samos and Kalymnos islands of Greece which abroad of our country, included the statistically analyses. As results of diagnosis, 4 new species belonging to Troglophilus and 1 new species belonging to Dolichopoda were found and preparing for publication. Within the molecular studies, sequence of 16S rDNA gene sequences are used (Caucasia, Crete, Samos and Kalymnos islands also included) for phylogenetical analyses (Bayesian analyses, molecular clock dating etc) and results are used for phylogenetical and biogeographical evaluations and hypothesis. Bu araştırmada, 2006-2010 yılları arasında Türkiye’nin farklı bölgelerinden toplanan Mağara Çekirgeleri (Rhaphidophoridae)’ne ait morfolojik ve moleküler veriler ışığında, mağara çekirgelerinin taksonomi, filogeni ve biyocoğrafyaları çalışılmıştır. Türkiye’de mağara çekirgeleri alanında yapılan ilk çalışma özelliğinde olan bu tezde, morfolojik çalışmalar kapsamında, Dolichopoda cinsine ait türlerden 19 ve Troglophilus cinsine ait 20 karaktere ait ölçümler alınmış ve mağara çekirgelerinin teşhislerinde kullanılan kalitatif karakterler incelenerek, ülkemizde bulunan mağara çekirgelerinin taksonomileri çalışılmıştır. Ayrıca ülkemiz dışında Kafkasya ve Yunanistan’ın Samos ve Kalymnos Adaları’ndan örneklere ait kalitatif ve kantitatif ölçümler alınıp analizlere dahil edilmiştir. Teşhisler sonucunda Troglophilus cinsine ait 4, Dolichopoda cinsine ait 1 yeni tür bulunmuş ve literatüre kazandırılmak üzere yayına hazırlanmaktadır. Moleküler çalışmalar kapsamında ise 16S rDNA geni sekansı kullanılarak (Kafkasya, Girit, Samos ve Kalymnos populasyonları dahil) filogenetik analizler (Bayesiyan Analizi, moleküler saat uygulamaları vb.) ve elde edilen sonuçlar ışığında filogenik ve biyocoğrafik değerlendirmeler ve varsayımlar yapılmıştır.
Book
Caves and other subterranean habitats with their often strange (even bizarre) inhabitants have long been objects of fascination, curiosity, and debate. The question of how such organisms have evolved, and the relative roles of natural selection and genetic drift, has engaged subterranean biologists for decades. Indeed, these studies continue to inform the general theory of adaptation and evolution. Subterranean ecosystems generally exhibit little or no primary productivity and, as extreme ecosystems, provide general insights into ecosystem function. The Biology of Caves and other Subterranean Habitats offers a concise but comprehensive introduction to cave ecology and evolution. Whilst there is an emphasis on biological processes occurring in these unique environments, conservation and management aspects are also considered. The monograph includes a global range of examples from more than 25 countries, and case studies from both caves and non-cave subterranean habitats; it also provides a clear explanation of specialized terms used by speleologists. This accessible text will appeal to researchers new to the field and to the many professional ecologists and conservation practitioners requiring a concise but authoritative overview. Its engaging style will also make it suitable for undergraduate and graduate students taking courses in cave and subterranean biology. Its more than 650 references, 150 of which are new since the first edition, provide many entry points to the research literature.
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