Campodea, Westwood, 1842

Sendra, Alberto & Reboleira, Ana Sofia P. S., 2020, Euro-Mediterranean fauna of Campodeinae (Campodeidae, Diplura), European Journal of Taxonomy 728, pp. 1-130 : 21-27

Subgenus Campodea s. str. Westwood, 1842

Campodea s. str. Westwood, 1842: 71.

Diagnosis

Notal formula with 1+1 medial anterior, 1+1 lateral anterior, 1+1 lateral posterior macrosetae on pronotum and mesonotum, 1+1 medial anterior, 1+1 lateral posterior on metanotum, besides a few species with 1+1 medial anterior, 1+1 lateral posterior or 1+1 lateral anterior, 1+1 lateral posterior on mesonotum and/or 1+1 lateral posterior or 0+0 macrosetae on metanotum; dorsal and lateral tarsal setae smooth; 0+0 or 1+1 lateral posterior macrosetae on IV–VII urotergites; from 0 to 1+1 lateral anterior macrosetae on IV–VII urotergites; 0+0 or 1+1 medial anterior or medial posterior macrosetae on I–IX urotergites; 3+3 lateral posterior macrosetae on VIII urotergite and 5+5 lateral posterior macrosetae on IX abdominal segment; first urosternite of the male with an area of continuous (in a few species slipt in two) glandular g 1 -setae (absent in a few species and in seasonal periods), subtrapezoidal appendages with glandular a 1 and a 2 -setae (a 2 -glandular setae absent in a few species); females without g 1 -glandular setae (with the exception of C. (C.) franzi ) with subcylindrical appendages with a

1

-glandular setae.

Habitat and distribution

The subgenus Campodea encompasses 92 described species spread across Europe and the Mediterranean Region, around the 60º parallel in the Scandinavian Peninsula ( Lie-Pettersen, 1898; Reuter 1895) to some spots of humid soils in North Africa ( Condé 1956a; Paclt 1957; Condé 1953b; Silvestri 1932b) ( Fig. 58 ). Only nine species colonize cave ecosystems distributed in karst regions on the East coast of the Iberian Peninsula and in the Western Mediterranean Islands ( Condé 1956a; Sendra & Moreno 2004). Most inhabit the soil and are well adapted to upper and lower soil horizons. The distribution of Campodea subgenus outside this region is poorly known but in the Middle East true Campodea have been observed in Iran ( Azadbakhsh & Nozari 2016). In Nearctic ecozone, Campodea subgenus is well distributed with a total of 13 species described in North America and Mexico ( Allen 1994, 1995; Condé 1956a; Condé & Geeraert 1962; Palacios-Vargas & García-Gómez 2014).

Taxonomic key for species of the subgenus Campodea s. str.

1. Lateral posterior metanotal macrosetae 1+1...................................................................................... 2 – Without lateral posterior metanotal macrosetae .............................................................................. 52

2. Medial posterior macrosetae of the ninth abdominal segment 1+1................................................... 3 – Without medial posterior macrosetae of the ninth abdominal segment........................................... 18

3. Lateral posterior macrosetae of the fifth urotergite 1+1 .................................................................... 4 – Without lateral posterior macrosetae of the fifth urotergite............................................................. 14

4. Medial posterior macrosetae of the seventh urotergite 1+1................................................................. .................................................................................... Campodea (Campodea) giardi Silvestri, 1912 – Without medial posterior macrosetae of the seventh urotergite ........................................................ 5

5. Medial posterior macrosetae of the eighth urotergite 1+1................................................................. 6 – Without medial posterior macrosetae of the eighth urotergite............................................................. ........................................................... Campodea (Campodea) consobrina Condé & Mathieu, 1957 6. Lateral posterior macrosetae of the fourth urotergite 1+1 ................................................................... ............................................................................ Campodea (Campodea) sardiniensis Bareth, 1980 – Without lateral posterior macrosetae of the fourth urotergite............................................................ 7

7. Medial anterior macrosetae of the second urotergite 1+1.................................................................. 8 – Without medial anterior macrosetae of the second urotergite ......................................................... 13

8. Cerci pilosity with thin macrosetae and smooth or with one to three apical tiny barbs......................

............................................................ Campodea (Campodea) sensillifera Condé & Mathieu, 1957 – Cerci pilosity with long poorly barbed macrosetae, shorter smooth clothing setae .......................... 9 – Cerci pilosity with latero interior swollen macrosetae in proximal articles .................................... 10 – Cerci pilosity of the C. plusiochaeta type ........................................................................................11

9. Male g 1 glandular setae present but separated; sensillum of the cupuliform organ simple; notal macrosetae short (medial anterior, lateral anterior) and long barbed lateral posterior; body shape not cave adapted........................................................ Campodea (Campodea) oredonensis Condé, 1951

– Male g 1 glandular setae present and continuous; sensillum of the cupuliform organ complex; notal macrosetae middle sized, robust, barbed; body shape cave adapted ................................................... .................................................................................... Campodea (Campodea) procera Condé, 1948

10. Male g 1 glandular setae present but separated; female without a 2 glandular setae; notal macrosetae short (medial anterior, lateral anterior) and long poorly barbed lateral posterior; marginal setae short smooth or with a few small distal barbs ...................... Campodea (Campodea) ilixonis Denis, 1932

– Male g 1 glandular setae present and continuous; female with a 2 glandular setae; notal macrosetae short (medial anterior, lateral anterior) and long barbed lateral posterior; marginal setae short with apical long barbs..................................................... Campodea (Campodea) meinerti Bagnall, 1918

11. Notal macrosetae long barbed; marginal setae long barbed ................................................................

................................................................................ Campodea (Campodea) zuluetai Silvestri, 1932 – Notal macrosetae long robust barbed; marginal setae long thin pine cone ..................................... 12 – Notal macrosetae short (medial anterior, lateral anterior) and long barbed lateral posterior; marginal

setae short with apical long barbs................................ Campodea (Campodea) lecreci Bareth, 1985

12. Medial anterior submacrosetae of the first urotergite 1+1; medial anterior submacrosetae of the second urotergite 1+1; medial anterior submacrosetae of the third urotergite 1+1; male without g 1 glandular setae .............................................................. Campodea (Campodea) egena Condé, 1951

– Without medial anterior submacrosetae of the first urotergite; without medial anterior submacrosetae of the second urotergite; without medial anterior submacrosetae of the third urotergite; male g 1 glandular setae present and continuous .. Campodea (Campodea) cossetana Sendra & Escolà, 2004

13. Male g 1 glandular setae absent; medial anterior submacrosetae of the first urotergite 1+1; medial anterior submacrosetae of the second urotergite 1+1; medial anterior submacrosetae of the third urotergite 1+1................................................................ Campodea (Campodea) egena Condé, 1951

– Male g 1 glandular setae present and continuous; without medial anterior submacrosetae of the first urotergite; wout medial anterior submacrosetae of the second urotergite; without medial anterior submacrosetae of the third urotergite................................................................................................... .................................................... Campodea (Campodea) maestrazgoensis Sendra & Escolà, 2004

14. Medial posterior macrosetae of the eighth urotergite 1+1.............................................................. 15 – Without medial posterior macrosetae of the eighth urotergite............................................................. ................................................................................ Campodea (Campodea) machadoi Condé, 1951

15. Lateral posterior metanotal short like marginal setae; sensillum of the cupuliform organ complex; one ventral sensilla on the third antennomere; body shape cave-adapted........................................... ............................................................ Campodea (Campodea) grallesiensis Sendra & Condé, 1987 – Lateral posterior metanotal long to middle sized; sensillum of the cupuliform organ simple; one dorsal sensilla on the third antennomere; body shape not cave adapted ......................................... 16

16. Lateral anterior macrosetae of the fifth urotergite 1+1; male g 1 glandular setae area present but separated in two; cerci pilosity with latero interior swollen macrosetae in proximal articles......... 17

– Without lateral anterior macrosetae of the fifth urotergite; male g 1 glandular setae present and continuous; cerci pilosity with long, barbed macrosetae and a few short smooth setae...................... ..................................................................... Campodea (Campodea) arrabidae Wygodzinsky, 1944

17. Notal macrosetae short (medial anterior, lateral anterior) and long (lateral posterior) robust barbed; marginal setae short with thin pine cone shape ....... Campodea (Campodea) rhopalota Denis, 1930

– Notal macrosetae middle sized, robust, poorly barbed; marginal setae with a long thin and pine cone shape ........................................................................... Campodea (Campodea) kervillei Denis, 1932

18 Lateral posterior macrosetae of the fifth urotergite 1+1 .................................................................. 19 – Without lateral posterior macrosetae of the fifth urotergite............................................................. 33

19. Lateral posterior macrosetae of the fourth urotergite 1+1 ............................................................... 20 – Without lateral posterior macrosetae of the fourth urotergite.......................................................... 24

20. Medial anterior macrosetae of the first to fourth urotergites 1+1.................................................... 21 – Without medial anterior macrosetae of the first to fourth urotergites.............................................. 23

21. Medial anterior macrosetae of the eighth urotergite 1+1; medial anterior macrosetae of the ninth abdominal segment 1+1; epicuticle with microdenticles..................................................................... ............................................................................. Campodea (Campodea) lankesteri Silvestri, 1912

– Without medial anterior macrosetae of the eighth urotergite; without medial anterior macrosetae of the ninth abdominal segment; epicuticle without microdenticles.................................................... 22

22. Lateral anterior macrosetae of the fourth to seventh urotergites 1+1.................................................. ....................................................................................... Campodea (Campodea) kasiki Rusek, 1964

– Without lateral anterior macrosetae of the fourth to seventh urotergites............................................. ..................................................... Campodea (Campodea) portacoeliensis Sendra & Jiménez, 1986

23. Lateral anterior macrosetae of the fourth urotergite 1+1; notal macrosetae short (medial anterior, lateral anterior) and long barbed lateral posterior; marginal setae short with apical long barbs......... ............................................................................... Campodea (Campodea) subdives Silvestri, 1933

– Without lateral anterior macrosetae of the fourth urotergite; notal macrosetae long thin barbed; marginal setae long barbed ........................................... Campodea (Campodea) redii Silvestri, 1912

24. Without swollen sensilla (Pagés’ setae) on labium.......................................................................... 25 – With swollen sensilla (Pagés’ setae) on labium...... Campodea (Campodea) chardardi Condé, 1947

25. Medial anterior macrosetae of the first to fourth urotergites 1+1.................................................... 26 – Without medial anterior macrosetae of the first fourth urotergites.................................................. 29

26. Lateral posterior metanotal short like marginal setae; lateral posterior mesonotal short like marinal setae; cerci pilosity with latero-interior swollen macrosetae in proximal articles............................... ........................................................................... Campodea (Campodea) minor Wygodzinsky, 1944

– Lateral posterior metanotal long to middle sized; lateral posterior mesonotal large to middle sized; cerci pilosity with long, barbed macrosetae and a few short smooth setae..................................... 27

27. Medial anterior macrosetae of the ninth urotergite 1+1; marginal setae short thick with a few barbs................................................................................................................................................. 28 – Without medial anterior macrosetae of the ninth urotergite; marginal setae short, smooth or with a few small distal barbs ............................................... Campodea (Campodea) epirotica Condé, 1984

28. Male g 1 glandular setae absent; 26–30 antennomeres; male a 2 glandular setae absent; notal macrosetae long, robust and barbed........................ Campodea (Campodea) delamarei Condé & Mathieu, 1957 – Male g 1 glandular setae present and continuous; 16–20 antennomeres; male a 2 glandular setae present; notal macrosetae long, robust and poorly barbed .................................................................. ............................................................................. Campodea (Campodea) frascajensis Condé, 1946

29. Lateral anterior macrosetae in the fifth urotergite 1+1 .................................................................... 30 – Without lateral anterior macrosetae in the fifth urotergite............................................................... 32

30. Clothing setae short thick with barbs or smooth; notal macrosetae long robust barbed; marginal setae short, thin and pine cone shape.......................... Campodea (Campodea) tuxeni Wygodzinsky, 1941

– Clothing setae long, thin and smooth; notal macrosetae long thin barbed; marginal setae long thin barbed ...................................................................... Campodea (Campodea) wallacei Bagnall, 1918

– Clothing setae short to middle sized, smooth or with a few small apical barbs; notal macrosetae short (medial anterior, lateral anterior) and long barbed lateral posterior; marginal setae short with apical long barbs......................................................................................................................................... 31

31. Cerci pilosity with long, barbed macrosetae and a few short smooth setae ........................................ ........................................................................ Campodea (Campodea) plusiochaeta Silvestri, 1912

– Cerci pilosity with short macrosetae and dense short smooth setae.................................................... ................................................................... Campodea (Campodea) pempturochaeta Silvestri, 1912

32. Lateral anterior macrosetae of the sixth and seventh urotergites 1+1; cerci pilosiy with long barbed macrosetae and a few short smooth setae; notal macrosetae short (medial anterior, lateral anterior), and long and barbed lateral posterior macrosetae................................................................................ ........................................................................ Campodea (Campodea) anae Sendra & Teruel, 2010

– Without lateral anterior macrosetae of the sixth and seventh urotergites; cerci pilosity with long, thin, smooth macrosetae; notal macrosetae long, robust and barbed .................................................. .................................................................................. Campodea (Campodea) ghilarovi Rusek, 1965

33. Medial anterior mesonotal and metanotal macrosetae 1+1 ............................................................ 34 – Without medial anterior mesonotal and metanotal macrosetae ...........................................................

..................................................................................... Campodea (Campodea) patrizii Condé, 1953

34. Female g 1 glandular setae absent ..................................................................................................... 35 – Female g

1

glandular setae present................................. Campodea (Campodea) franzi Condé, 1954

35. Labium type Pagés’ sensilla absent ................................................................................................. 36 – Labium type Pagés’ sensilla present ....................... Campodea (Campodea) chardardi Condé, 1947

36. Without lateral posterior metanotal macrosetae ..................................................................................

.................................................................... Campodea (Campodea) coniphora Wygodzinsky, 1941 – Lateral posterior metanotal short like marginal setae...................................................................... 37 – Lateral posterior metanotal middle sized to long ............................................................................ 42

37. Medial anterior macrosetae of the first urotergite 1+1; medial anterior macrosetae of the second urotergite 1+1; medial anterior macrosetae of the third urotergite 1+1; medial anterior macrosetae of the fourth urotergite 1+1.................................................................................................................. 38

– Without medial anterior macrosetae of the first urotergite; without medial anterior macrosetae of the second urotergite; without medial anterior macrosetae of the third urotergite; without medial anterior macrosetae of the fourth urotergite .................................................................................................. 40

38. Lateral posterior mesonotal as short as marginal setae.................................................................... 39 – Lateral posterior mesonotal middle sized to long................................................................................ ........................................ Campodea (Campodea) machadoi Condé, 1951 (specimens from Braga)

39. Clothing setae short and thick with barbs or smooth; with short macrosetae and abundant short, smooth setae; notal macrosetae short, thick and poorly barbed; marginal setae short thick with a few barbs...................................................................... Campodea (Campodea) lubbockii Silvestri, 1912 – Clothing setae short to middle sized, smooth or with a few small apical barbs; cerci pilosity with latero-interior swollen macrosetae in proximal articles; notal macrosetae short (medial anterior, lateral anterior) and long barbed lateral posterior; marginal setae short, smooth or with a few small distal barbs......................................................... Campodea (Campodea) minor Wygodzinsky, 1944

40. Cerci pilosity with long and poorly barbed macrosetae, shorter smooth clothing setae ................ 41 – Cerci covered with whorls of thin macrosetae bent forward and backward, and surrounded by a few clothing setae ...................................................... Campodea (Campodea) westwoodi Bagnall, 1918

41. Lateral anterior macrosetae of the fifth urotergite 1+1; smooth clothing setae short to middle sized or with a few small apical barbs; epicuticle without microdenticles; trochanteral sensillum none..... .................................................................... Campodea (Campodea) coniphora Wygodzinsky, 1941

– Without lateral anterior macrosetae of the fifth urotergite; clothing setae short and thick with barbs or smooth; epicuticle with microdenticles; trochanteral sensillum one............................................... .................................................................................. Campodea (Campodea) grassii Silvestri, 1912

42. Lateral posterior macrosetae of the sixth and seventh urotergites 1+1........................................... 43 – Without lateral posterior macrosetae of the sixth and seventh urotergites ..........................................

.............................................................................. Campodea (Campodea) vihorlatensis Paclt, 1961

43. Medial anterior macrosetae of the first to fourth urotergites 1+1........................................................ .................................................................... Campodea (Campodea) basiliensis Wygodzinsky, 1940 – Without medial anterior macrosetae of the first to fourth urotergites.............................................. 44

44. Cerci pilosity long and poorly barbed macrosetae; shorter, smooth clothing setae......................... 45

– Cerci bearing one complete whorl of short macrosetae with one apical barb and numerous short clothing setae for each article .......................... Campodea (Campodea) pseudofragilis Condé, 1984

– Cerci pilosity with latero interior swollen macrosetae in proximal articles ........................................ ................................................................................... Campodea (Campodea) taunica Marten, 1939

– Cerci pilosity with long and barbed macrosetae and a few short smooth setae .............................. 48

– Cerci pilosity with short macrosetae and abundant short smooth setae .............................................. .................................................................................. Campodea (Campodea) fragilis Meinert, 1865

45. Lateral anterior macrosetae of the sixth and seventh urotergites 1+1 ............................................. 46 – Without lateral anterior macrosetae of the sixth and seventh urotergites............................................

................................................................................ Campodea (Campodea) pieltaini Silvestri, 1932

46. Lateral anterior macrosetae of the fifth urotergite 1+1; epicuticle without microdenticles................. .................................................................... Campodea (Campodea) coniphora Wygodzinsky, 1941 – Without lateral anterior macrosetae of the fifth urotergite; epicuticle with microdenticles ............ 47

47. Clothing setae thin pine cone shape; notal macrosetae middle sized, robust, barbed; marginal setae long thin pine cone shape; without trochanteral sensillum.................................................................. ....................................................................... Campodea (Campodea) monspessulana Condé, 1953

– Clothing setae short to middle sized, smooth or with a few small apical barbs; notal macrosetae middle sized, robust and poorly barbed; marginal setae short thin pine cone shape; one trochanteral sensillum.................................................................... Campodea (Campodea) magna Ionescu, 1955

48. Lateral anterior macrosetae of the fifth urotergite 1+1 .................................................................... 49 – Without lateral anterior macrosetae of the fifth urotergite............................................................... 50

49. Male a 2 glandular setae absent; notal macrosetae short (medial anterior, lateral anterior) and long barbed lateral posterior; marginal setae short with apical long barbs.................................................. ................................................................................ Campodea (Campodea) silvestrii Bagnall, 1918

– Male a 2 glandular setae present; notal macrosetae long, thin, smooth or poorly barbed; marginal setae short, smooth or with a few small distal barbs ........................................................................... ....................................................................... Campodea (Campodea) silvicola Wygodzinsky, 1940

50. Lateral anterior macrosetae of the sixth urotergite 1+1; lateral anterior macrosetae of the seventh urotergite 1+1................................................................................................................................... 51

– Without lateral anterior macrosetae of the sixth urotergite; without lateral anterior macrosetae of the seventh urotergite........................................... Campodea (Campodea) sarae Sendra & Teruel, 2010

51. Clothing setae thin with pine cone shape ...... Campodea (Campodea) monspessulana Condé, 1953 – Clothing setae short with apical barbs...................... Campodea (Campodea) augens Silvestri, 1936 – Clothing setae short to middle sized, smooth or with a few small apical barbs..................................

............................................................................... Campodea (Campodea) posterior Silvestri, 1932

52. Lateral posterior macrosetae of the fifth urotergite 1+1 .................................................................. 53 – Without lateral posterior macrosetae of the fifth urotergite............................................................. 55

53. Lateral anterior macrosetae of the sixth and seventh urotergites 1+1; cerci pilosity with latero interior swollen macrosetae in proximal articles.......................................................................................... 54

– Without lateral anterior macrosetae of the sixth and seventh urotergites; cerci pilosity with long, barbed macrosetae and a few short smooth setae ... Campodea (Campodea) alluvialis Sendra, 2017

54. Lateral anterior macrosetae of the fifth urotergite 1+1; male g 1 glandular setae absent...................... ........................................................................ Campodea (Campodea) galilaea Wygodzinsky, 1942 – Without lateral anterior macrosetae of the fifth urotergite; male g 1 glandular setae present and continuous.............................................................. Campodea (Campodea) galilaea (Sardinia form)

55. Medial anterior mesonotal macrosetae 1+1; medial anterior metanotal macrosetae 1+1 .............. 56 – Without medial anterior mesonotal macrosetae; without medial anterior metanotal macrosetae .......

........................................................................................ Campodea (Campodea) remyi Denis, 1930

56. Lateral anterior mesonotal macrosetae 1+1..................................................................................... 57 – Without lateral anterior mesonotal macrosetae.. Campodea (Campodea) apennina Ramellini, 1998

57. Lateral posterior macrosetae of the sixth urotergite 1+1 ................................................................. 58 – Without lateral posterior macrosetae of the sixth urotergite................................................................

.................................................................................. Campodea (Campodea) majorica Condé, 1955

58. Lateral anterior macrosetae of the fifth urotergite 1+1 ........................................................................ ................................................................................ Campodea (Campodea) blandinae Condé, 1948 Without lateral anterior macrosetae of the fifth urotergite............................................................... 59

59. Lateral anterior macrosetae of the sixth urotergite 1+1; lateral anterior macrosetae of the seventh urotergite 1+1................................................................................................................................... 60

– Without lateral anterior macrosetae of the sixth urotergite; without lateral anterior macrosetae of the seventh urotergite....................................... Campodea (Campodea) navasi Sendra & Moreno, 2004

60. Male g 1 glandular setae absent; cerci pilosity with long barbed macrosetae and a few short smooth setae; notal macrosetae middle size, barbed; marginal setae short thin pine cone shape.................... ..................................................................................... Campodea (Campodea) cyrnea Condé, 1946 – Male g 1 glandular setae present and continuous; cerci pilosity with long poorly barbed macrosetae and short smooth setae; notal macrosetae short (medial anterior, lateral anterior) and long barbed lateral posterior; marginal setae short with apical long barbs ............................................................. ....................................................................... Campodea (Campodea) staphylinus Westwood, 1842

Figures

Figs 58–59. Distribution in the Euro-Mediterranean region, includingAzores and Canaries.58. Subgenus Campodea s. str. Silvestri, 1932. 59. Subgenus Dicampa Silvestri, 1932.