Life in the Puddle

Darwin’s Warm Little Pond

The following might have been a better start to our 2001 paper (Athol McLachlan and Richard Ladle 2001. Life in the puddle: behavioural and life-cycle adaptation in the Diptera of tropical rain pools Biol. Rev., 76, 377-388).

ABSTRACT

For over one hundred years Charles Darwin’s “warm little pond” has been at the centre of conjecture about the origin of life. It is precisely such ponds that form the subject of the present article. These tropical ponds have some extraordinary properties. Among them is the fact that they harbour exceedingly high densities of single species of midge larvae, each carrying the clear stamp of adaptation to the durational characteristics of their pond. Exceptional too, is the property which derives from their occupying depressions on rock surfaces, of being spatially consistent over millions of years. In what follows we attempt to draw out these properties and set them against those of other very transient habitats.

INTRODUCTION

Puddles of rain water on the surfaces of rock exposures are a little known but very common habitat for freshwater-dwelling animals. They are also typical of Charles Darwin’s “warm little pond” (Conway Morris, S. 2003, p53, 64, Dawkins, R. 2009, p417, 419). In Africa, these are inhabited by the larvae of two taxa of fly unique to these pools. One of these includes species able to survive dry periods in situ; the other includes species that must reach adulthood and migrate to survive periods when the pool is dry. Hence, the opportunity exists for a comparative study of adaptation among these species. Since puddles are small, our principal method in the study of adaptation has been the experimental manipulation of puddles and their faunas in the wild. Using this method we were able to identify the spatial consistence of pools and their unpredictable duration during the rainy season as the main selective pressures shaping adaptation. Adaptations include diapause and adaptive adjustments of the life cycle. It is the second of these that provides the focus of our interest here. There are many kinds of freshwater habitat ……run on to original version (Introduction line 1).

Refrences

Darwin, C. 1871. The Descent of Man and Selection in Relation to Sex. J. Murray, London.

Conway Morris, S, 2003. Life’s Solutions: Inevitable Humans in a Lonely Universe. Cambridge University Press, Cambridge, UK.

Dawkins, R. 2009. The Greatest Show on Earth. Bantam Press, London.

My Published Work in the 'learned literature' 1965-2011

Theses, Published Symposium Proceedings and Reports

1965. Ecology of the bottom fauna of Lake Kariba, pp. 58- 59. In: LKFRI Kariba Research Symposium Proceedings.

1968. A study of the bottom fauna of Lake Kariba. PhD thesis, University of London. 348 pp.

1969. Aspects of the ecology of the bottom fauna of Lake Chilwa, pp. 27-29. In: IBP Symposium, Malawi Proceedings.

1969. A study of the bottom fauna of Lake Kariba, pp. 13-14. In: Nuffield Lake Kariba Research Sation Report, 1962-68. University College of Rhodesia Press, Salisbury.

1974. The development of chironomid communities in a new temperate impoundment. Ent.Tidskr., 95 (Suppl.): 162-71.

1978. Interactions between freshwater animals and micro-organisms. Ann.appl.Biol., 89: 162-65.

1980. Chironomids and particles: Micro-organisms and chironomid distribution in a peaty upland river. In: D A Murray (ed) Chironomidae: Ecology, Systematics, Cytology and Physiology. pp. 171-77. Pergamon Press, Oxford (with R J Toscano).

1980. Chironomids and particles: A field experiment with peat in an upland stream. In: D A Murray (ed) Chironomidae: Ecology, Systematics, Cytology and Physiology. pp. 179-85. Pergamon Press, Oxford (with A T Walentowicz).

1981.Biological consequences of fluctuations in lake level. In: J M Kapetsky (ed) Seminar on river basin development (IFA Technical report No 8) pp. 225-31.

Peer reviewed original research papers

1. McLachlan AJ (1969). Notes on some larval and pupal chironomids (Diptera) from Lake Kariba,Rhodesia J.Nat.Hist., 3: 261- 93.

2. McLachlan AJ (1969). Substrqate preferences and invasion behaviour exhibited by larvae of Nilodorum brevibucca Freeman (Chironomidae) under experimental conditions. Hydrobiologia, 33: 237-249.

3. McLachlan AJ (1969). The effect of aquatic macrophytes on the variety and abundance of benthic fauna in a newly created lake in the tropics (Lake Kariba). Arch. Hydrobiologia, 66: 212-31.

4. McLachlan AJ and McLachlan SM (1969). The bottom fauna and sediments in a drying place of a saline African lake (L. Chilwa, Malawi). Hydrobiologia, 34: 401-13.

5. McLachlan AJ (1970). Some effects of annual fluctuations in water level on the larval chironomid communities of Lake Kariba. J.Anim.Ecol., 39: 79-90.

6. McLachlan AJ (1970). Submerged trees as a substrate for benthic fauna in the recently created Lake Kariba (Central Africa). J.Appl.Ecol., 7: 253-66.

7. McLachlan AJ (1971). Some immature stages of the subgenus Chironomus (Meigen) (Diptera: Chironomidae) from Malawi, Central Africa. J.Ent.(B), 40: 173-78.

8. McLachlan AJ and McLachlan SM (1971). Benthic fauna and sediments in the newly created Lake Kariba (Central Africa). Ecology, 52: 800-09.

9. McLachlan AJ, Morgan PR, Howard-Willians C, McLachlan SM and Bourn D (1972). Aspects of the recovery of a saline African lake following a dry period. Arch.Hydrobiol., 70: 325-40.

10. McLachlan AJ (1974). Recovery of the mud substrate and its associated fauna following a dry phase in a tropical lake. Limnol.Oceanogr., 19: 74-83.

11. McLachlan AJ (1975). The role of aquatic macrophytes in the recovery of the benthic fauna of a tropical lake after a dry phase. Limnol. Oceanogr., 20: 54-63.

12 McLachlan AJ and McLachlan SM (1975). The physical environment and bottom fauna of a bog lake. Arch.Hydrobiol., 76: 198-217.

13. McLachlan AJ (1976). Factors restricting the range of Glyptotendipes paripes Edwards (Diptera: Chironomidae) in a bog lake. J.Anim.Ecol., 45: 105-113.

14. McLachlan AJ amd McLachlan SM (1976). Development of the mud habitat during the filling of two new lakes. Freshwat.Biol., 6: 59-67.

15. Mclachlan AJ (1976). Variation in ‘gill’ size in a larva of the African midge Chironomus transvaalensis Keiffer. Limnol.Soc.Stn.Afr., 2: 55- 56.

16. McLachlan AJ and Cantrell MA (1976). Sediment development and its influence on the distribution and tube structure of Chironomus plumosus L. (Chironomidae: Diptera) in a new impoundment. Freshwat.Biol., 6: 437-43.

17. McLachlan AJ (1977). Some effects of tube shape on the feeding of Chironomus plumosus L. (Diptera: Chironomidae). J.Anim.Ecol. 46: 139-46.

18. McLachlan AJ (1977). The changing role of terrestrial and autochthonous organic matter in newly flooded lakes. Hydrobiologia, 54: 215- 17.

19. McLachlan AJ (1977). Density and distribution in laboratory populations of midge larvae (Chironomidae: Diptera). Hydrobiologia, 55: 195- 99.

20. Cantrell MA and McLachlan AJ (1977). Competition and chironomid distribution patterns in a newly flooded lake. Oikos, 29: 429-33.

21. McLachlan AJ and Dickinson CH (1977) Micro-organisms as a factor in the distribution of Chironomus lugubris in a bog lake. Archiv fur Hyrobiology, 80: 133-146.

22. Brennan A, McLachlan AJ & Wotton RS (1978). Particulate material and midge larvae (Chironomidae:Diptera) in an upland river. Hydrobiologia, 59: 67-73.

23. McLachlan AJ, Brennan A andWotton RS (1978). Particle size and chironomid (Diptera) food in an upland river. Oikos, 31: 247-52.

24. Baker A and McLachlan AJ (1979). Food preferences of Tanypodinae larvae (Diptera: Chironomidae). Hydrobiologia, 62: 283-88.

25. Brennan A and McLachlan AJ (1979). Tubes and tube building in a lotic chironomid (Dipteran) community. Hydrobiologia, 67: 173-78.

26. McLachlan AJ, Pearce LJ and Smith JA (1979). Feeding interactions and cycling of peat in a bog lake. J.Anim.Ecol., 48: 851-61.

27. Brennan A and McLachlan AJ (1980). Species of Eukieffreriella Theinemann (Diptera: Chironomidae) from a northern river, with some notes on larval dwellings. Ent.Mon.Mag.16: 109-11.

28. McLachlan AJ and Cantrell MA (1980). Survival strategies in tropical rain-pools. Oecologia Berl., 47: 344-51.

29. McLachlan AJ (1981). Food sources and foraging tactics in tropical rain-pools. Z.J.Linn.Soc. 75: 267-277.

30. McLachlan AJ (1981). Interactions between insect larvae and tadpoles in tropical rain-pools. Ecol.Ent., 6: 175-82.

31. Cantrell MA and McLachlan AJ (1982). Habitat duration and dipteran larvae in tropical rain-pools. Oikos, 38: 343-48.

32. McLachlan AJ (1983). Life history tactics of rain-pool dwellers. J.Anim.Ecol. 52: 545-61.

33. McLachlan AJ (1983). Habitat distribution and body size in rain-pool dwellers. Zool.J.Linn.Soc., 79: 399-407.

34. McLachlan AJ (1985). The relationship between habitat predictability and wing length in midges (Chironomidae). Oikos, 44: 391-97.

35. McLachlan AJ (1985). What determines the species present in a rain-pool? Oikos, 45: 1-7.

36. Osborne PL andMcLachlan AJ (1985). The effect of tadpoles on algal growth in temporary, rain- filled rock pools. Freshwat.Biol., 15: 77-87.

37. McLachlan AJ (1986). Sexual dimorphism in midges; strategies for flight in the rain-pool dweller Chironomus imicola (Diptera: Chironomidae). J.Anim.Ecol., 55: 261-67.

38. Burtt ET, Perry RJO and McLachlan AJ (1986). Feeding and sexual dimorphism in adult midges (Diptera:Chironomidae). Holarctic Ecology, 9: 27-32.

39. McLachlan AJ (1986). Survival of the smallest: advantages and costs of small size in flying animals. Ecol.Ent.,11: 237-40.

40. McLachlan AJ and Allen DA (1987). Male mating success in Diptera: advantages of small size. Oikos, 48: 11-14

41. McLachlan AJ (1988). Refugia and habitat partitioning among midges (Diptera: Chironomidae) in rain-pools. Ecol.Ent., 13: 185-93.

42. McLachlan AJ and Neems RM (1989). An alternative mating system in small male insects. Ecol.Ent., 14: 85-91.

43. McLachlan AJ and Yonow T (1989). Reproductive strategies in rain-pool dwellers and the model freshwater insect. Hydrobiologia, 171: 223-230.

44. Patterson JW and McLachlan AJ (1989). Larval habitat duration and size at metamorphosis in frogs. Hydrobiologia, 171: 121- 126.

45. Neems RM, McLachlan AJ and Chambers R (1990). Body size and lifetime mating success of male midges (Diptera:Chironomidae). Animal Behaviour., 40: 648-652.

46. Jackson JM and McLachlan AJ (1991). Rain-pools on peat moorland as island habitats for midge larvae. Hydrobiologia, 209: 59-67.

47. Neems RM, Lazarus J and McLachlan AJ (1992). Swarming behaviour in male chironomid midges; a cost-benefit analysis. Behavioural Ecology, 3: 285-290.

48. McLachlan AJ and Neems RM (1993). Are females selected against in ephemeral habitats? The case of a holometabolous insect (Chironomus pulcher). Oecologia, 94: 83-86.

49. McLachlan AJ and Cant M (1995). Small males are more symmetrical: mating success in the midge Chironomus plumosus (Diptera:Chironomidae). Animal Behaviour, 50: 841-846.

50. McLachlan AJ and Neems RM (1996). Is flight architecture determined by physical constraints or by natural selection: the case of the midge Chironomus plumosus. Journal of Zoology (London), 240: 301-308.

51. McLachlan AJ (1997). Size or symmetry: an experiment to determine which of the two accounts for mating success in male midges. Ecoscience: 4: 454-459.

52. Neems RM, Lazarus J and McLachlan AJ (1998). Lifetime reproductive success in a swarming midge: Trade-offs and stabilizing selection for male body size. Behavioural Ecology, 9: 249-256.

53. McLachlan AJ (1999). Parasites promote mating success: the case of a midge and a mite. Animal Behaviour, 57: 1199-1205.

54. McLachlan AJ, Ladle R and Bleay C (1999). Is infestation the result of adaptive choice behaviour by the parasite? A study of mites and midges. Animal Behaviour, 58: 615-620.

55. McLachlan AJ, Ladle R and Crompton B (2003). Predator-prey interactions on the wing: aerobatics and body size among dance flies and midges. Animal Behaviour, 66: 911-915.

56. Crompton B, Thomason J and McLachlan A (2003). Mating in a viscous universe: the race is to the agile, not to the swift. Proceedings of the Royal Society, London (B), 270:1991-1995.

57. McLachlan AJ (2006). You are looking mitey fine: parasites as direct indicators of fitness in the mating system of a host species. Ethology Ecology & Evolution, 18: 233-239.

58. McLachlan AJ, Pike, TW and Thomason JC (2008). Another kind of symmetry: are there adaptive benefits to the arrangement of mites on an insect host? Ethology Ecology and Evolution, 20: 257-270.

Reviews, Debates and Book Chapters

1. McLachlan AJ (1969). Some effects of water level fluctuation on the benthic fauna of two central African lakes. Limnol.Soc.S.Afr.Newsl., 13: 58-63.

2. McLachlan AJ (1970). The bottom fauna: distribution patterns and invasion behaviour. pp. 34-35. In: Kalk, M (ed) Decline and recovery of a saline Lake. Lake Chilwa Coordinated Research Report, 1966- 70. Government Printer, Zomba (with S M McLachlan).

3. McLachlan AJ (1974). Development of some lake ecosystems in Tropical Africa with special reference to the invertebrates. Biol.Rev., 49: 365-97.

4. McLachlan AJ (1979). Decline and recovery of the benthic invertebrate communities. pp. 143-60. In: Kalk et al (eds.) Lake Chilwa. Studies of change in a tropical ecosystem. Monographiae biologicae, 35. Junk. The Hague.

5. McLachlan AJ (1979). The aquatic environment: I. Chemical and physical characteristics of Lake Chilwa.pp.59-78. In: Kalk et al. (eds.) Lake Chilwa. Studies of change in a tropical ecosystem.

6. McLachlan AJ (1986). Chironomid wing length: a measure of habitat duration and predictability? A reply to Vepsalainen. Oikos, 46: 271-73.

7. McLachlan AJ. (1988). Male mating success in Diptera: A reply to Thompson. Oikos, 51: 109.

8. McLachlan AJ (1988). Animal populations at extreme densities: size dimorphism by frequency dependent selection, in ephemeral habitats. Functional Ecology, 3: 633-43.

9. McLachlan AJ and Neems RM. (1995). Swarm-Based Mating Systems. In: Insect Reproduction (eds SR Leather & J Hardie). CRC press, London.

10. McLachalan AJ and Ladle R. (2001). Life in the puddle: behavioural and life-cycle adaptations in the Diptera of tropical rain pools. Biological Reviews. 76: 377-388.

11. McLachlan AJ and Ladle R. (2008). The evolutionary ecology of detritus feeding in the larvae of freshwater Diptera. Biological Reviews, 84: 133-141.

12. McLachlan, AJ and Ladle, R (2010). Barriers to Adaptive Reasoning in Community Ecology. Biological Reviews, 86, 543-548. (doi: 10.1111/j.1469-185X.2190.00159.x).

13. McLachlan, AJ and Ladle, R (2010). Fluctuating Asymmetry in Flies, What Does It Mean? Symmetry, 2, doi:10.3390/sym2021099, 1099-1107.13.

14. McLachlan, AJ (2011). Homosexual Pairing within a Swarm-Based mating System: The Case of the Chironomid Midge. Article ID 854820, 5 pages (doi: 10.1155/2011/854820).