Sphecid and Carbronid wasps construct the characteristic nests with unique architectural patterns to lay their eggs and nurture their larvae. The present investigation was carried out the nest characteristic features and prey selection of mud dauber wasp from the A.V.C. College campus from January to March 2019. It deals with nest characteristic features of the active and abandoned nest of mud dauber wasp, studies indicate that Sceliphron madraspatanum nests are built on bright surfaces that are covered against water and immediate sunlight. The nests' height from the ground was less than 2 feet, nest length was less than 5 cm and cell depth or width was less than 3 cm. The mean number of orifices and cells was 6.0 and the cell length range was measured from 2.25 to 1.92 cm. Nests' shape was a maximum rectangle and a few were irregular shapes. In active nests, mean height from the ground and orifices or cells were lesser than the inactive nests. Likewise, the nest length and depth of the cells or width, weight were higher than the inactive nests. A total of 24 larvae were isolated from the 20 cells and weighed around 3.15 g. Similarly, the feed was also isolated and tally as 153 paralyzed individuals which belong to order Araneae and weighed 17.27 g. It concluded that the nest of mud dauber wasp showed species-specific architecture and it feed the spiders of an agroecosystem. From this research, the wasp might be considered a beneficial insect for the agroecosystem.
Nest Characteristic Features and Prey Selection of Mud Dauber Wasp Sceliphron madraspatanum “Fabricius, 1781”
Paramanandham Joothi1*, Revathi Arunagiri1, Sankari Ambalavanan1, Jayakumar Samidurai1, Krishnappa Kaliyamoorthy1, Ronald Ross Pankirias2
1PG and Research Department of Zoology and Wildlife Biology, A.V.C. College (Autonomous), Mannampandal- 609 305, Mayiladuthurai, Tamil Nadu, India.
2Department of Zoology, Govt. Arts College, Ariyalur- 621 713, Tamil Nadu, India.
ABSTRACT
Sphecid and Carbronid wasps construct the characteristic nests with unique architectural patterns to lay their eggs and nurture their larvae. The present investigation was carried out the nest characteristic features and prey selection of mud dauber wasp from the A.V.C. College campus from January to March 2019. It deals with nest characteristic features of the active and abandoned nest of mud dauber wasp, studies indicate that Sceliphron madraspatanum nests are built on bright surfaces that are covered against water and immediate sunlight. The nests' height from the ground was less than 2 feet, nest length was less than 5 cm and cell depth or width was less than 3 cm. The mean number of orifices and cells was 6.0 and the cell length range was measured from 2.25 to 1.92 cm. Nests' shape was a maximum rectangle and a few were irregular shapes. In active nests, mean height from the ground and orifices or cells were lesser than the inactive nests. Likewise, the nest length and depth of the cells or width, weight were higher than the inactive nests. A total of 24 larvae were isolated from the 20 cells and weighed around 3.15 g. Similarly, the feed was also isolated and tally as 153 paralyzed individuals which belong to order Araneae and weighed 17.27 g. It concluded that the nest of mud dauber wasp showed species-specific architecture and it feed the spiders of an agroecosystem. From this research, the wasp might be considered a beneficial insect for the agroecosystem.
Keywords: Mud dauber wasp, Nest architecture, Nest orientation, Prey, Spider, Paralyzed.
INTRODUCTION
Wasps are social or solitary nest builders by natural or artificial materials as a substrate. There are 35 species described in the genus Sceliphron worldwide [1]. A few definite groups of sphecid and crabronid wasps construct the characteristic nests with unique architectural patterns to lay their eggs and nurture their larvae. These nests are built out of mud unruffled from nearby mud puddles and hence these wasps are popularly nominated as the “mud daubers” or even the “dirt daubers”. Sphecidae is a cosmopolitan family of bees with solitary or altruistic demeanors that contains digger wasps, for example, mud bees, Sceliphron madraspatanum (Fabricius), and other familiar species, all of which fall into the category of thread-waisted wasps. Nest construction of wasps is very important to protect their larvae from enemies or predators and store the chow for long life. Specific architectural ability and materials used for nest construction are species-specific in the group of wasps. Earlier, few studies have been done in various species of wasps in the aspects of nest characteristic features and prey selection [2-5].
The individual forage search profession defines a circumstance in which homogeneous individuals in a general population show dissimilarities in food search behavior, each of which specializes in various kinds of prey. Individual expertise is widespread in animals but has not been studied in invertebrates, despite its potential effects on the food networks and population dynamics [6]. In the genus Sceliphron, females create their mud nests in various isolated and dry places, but some of them are usually related to human residential. A single cellule is first created, prepared with a spider, and sealed by a female before working on another cell. Typically, more than one cellule is created to form a cumulative nest. Cells that are not fully accumulated in a day may be temporarily sealed overnight [7, 8].
Generally, many cells of a nest are occult with a mud layer [9-12]. Halder et al. [2] have been observed to make around 10–13 round trips to construct each cell (2.38±0.19 cm x 0.975±0.083 cm) carrying soft mud collected mainly from frequently irrigated fields or nearby irrigation channels with their mandibles and forelegs in S. madraspatanum.
This study was carried out the nest architectural characteristics features and site selection for the nest construction. Moreover, the present study also deals with the prey availability in the nest cells of active nests of Mud dauber wasps.
MATERIALS AND METHODS
The nests were obtained from A.V.C. College Campus, Mannampandal, Mayiladuthurai, Tamil Nadu. The study area belongs to the vegetable agroecosystem and cultivated several vegetable crops like brinjal, bhendi, banana, etc., A total of 25 nests of S. madraspatanum were collected randomly during the period from January to March 2019 every week (every week two to three nests were collected and analyzed) and taken to the laboratory and put in plastic boxes (10 x 8 x 5 cm). Directly after opening the nests to analyze the cells and their structure. Bait was separated from the cells in terms of species and counted manually. The designation of spiders has been done using the thematic expertise and textbook of Indian spiders [13]. Some direct comments of nests were made at the nest site. The statistical analyses are diversity indices and correlation matrix proved the significance of the study and it was done by using IBM SPSS version 25.
RESULTS AND DISCUSSION
Sceliphron madraspatanum mud nests are well lit on surfaces and covered from water and direct sunlight. Some of them were huge, with a solid base, and were challenging to separate. Mud dauber nests showed that mean height from the ground was 52.44 ± 28.86 cm and the average length and width were 4.78±1.67 and 2.35±0.44 cm respectively. The weight of the nest was also studied, the mean weight was 13.94±11.70 g. The mean number of orifices was showed 6.0±1.58 and recorded the highest 10 orifices and four orifices were least. The number of chambers in the nest was 6.0±1.58 (n=25) and the range of the chamber or cell length was 2.25 to 1.91cm. Nests' shape was maximum rectangle (96%) and few were irregular (4%) and nest substrate orientation was 40% east direction followed by 32% north and 28% west direction. The nature of nest substrate was nearly 80% were walls followed by all other materials have each 4% namely, bamboo poles, strips, cloth, switch box and tables. The nest status also noted its shows nearly 84% of the nest were inactive, with the rest of them being active and were taken further studies such as feed on the chambers.
Among the 25 nests of Mud daubers, four nests were found as active nests and it was showed the mean height from the ground was 32±14.99 cm. Likewise, the nest length and width average was noted 5.13±1.45 and 2.48±0.21 respectively. The nest varied among the four nests from 10.26 g to 42.49 g and the average was calculated as 21.94±7.50 g. The number of orifices also varied according to their size and weight, it shows four to six and a mean calculated as 5±0.41.
Active nests showed that the mean number of chambers was 5.0±0.7 and length range between 2.25 to 1.91cm. The chamber diameter range was 0.52 to 0.46cm. Active nests were opened for analyzing the prey and larvae of Mud dauber, a total of 24 larvae were isolated from the four active nests and weighed around 3.15 g. Similarly, the feed was also isolated and tally as 153 individuals which belong to order Araneae and weighed 17.27 g. Among the four active nests of Mud dauber, two nests alone have been noticed that the prey items of family Salticidae (5 genera) were showed the highest frequency as well as diversity followed by Lycosidae (1 genus) and Araneidae (1 genus) (Table 1).
Table 1. Spider recorded inactive nests of S. madraspatanum during the study period (n=2)
Species of Spiders |
Common name |
Family |
Nest I |
Nest II |
Hyllus semicupreus |
Semi-coppered heavy jumper |
Salticidae |
20 |
17 |
Hasarius adansoni |
Adanson's house jumper |
1 |
1 |
|
Menemerus bivittatus |
Grey Wall jumper |
1 |
1 |
|
Bavia kairali |
Jumping spider |
41 |
13 |
|
Telamonia dimidiata |
Two striped Jumper |
9 |
1 |
|
Lycosa Mackenziei |
Wolf spider |
7 |
3 |
|
Neoscona Nautica |
Brown sailer spider |
6 |
4 |
|
Un identification |
|
|
19 |
9 |
Total |
|
|
104 |
49 |
The diversity indices showed, there is not much variation among the two nests of Mud daubers. The species richness, dominance index, Simpson diversity index, Shannan’H index, and Evenness are quite the same and individuals vary among the nests (Table 2).
The active nest shape of the mud daubers was rectangle shape and the substrate orientation of the nest shows 75% west followed by 25% east orientation. The substrate of the nest was preferred as the wall. Statically proved that the water sources positively influence the other dependent factors like nest height from the ground, nest length, number of orifices, and waste materials. The nest height from the ground positively influenced nest length alone and nest length positively influenced the nest weight. The nest length significantly influenced the number of orifices and nest weight (Table 3). The correlation matrix for active nests indicated that nest height from the ground significantly influenced spider weight, number of larva in the nest, and larval weight (Table 4).
Table 2. Diversity indices of spiders inactive nest of Mud dauber (n=2)
Indices |
Nest I |
Nest II |
Species richness |
8 |
8 |
Individuals |
104 |
49 |
Dominance Index |
0.2413 |
0.2362 |
Simpson Diversity index |
0.7587 |
0.7638 |
Shannon H' Index |
1.642 |
1.644 |
Evenness |
0.6456 |
0.6472 |
Table 3. Correlation matrix of variables irrespective of nests S. madraspadnum collected during the study period
|
Water source |
Nest height |
Nest width |
Nest length |
No. of orifices |
Nest weight |
Waste materials |
|
Water source |
Pearson Correlation Sig. (2-tailed) N |
1
25 |
|
|
|
|
|
|
Nest height |
Pearson Correlation Sig. (2-tailed) N |
0.226 0.277 25 |
1
25 |
|
|
|
|
|
Nest width |
Pearson Correlation Sig. (2-tailed) N |
-0.113 0.591 25 |
-0.192 0.357 25 |
1
25 |
|
|
|
|
Nest length |
Pearson Correlation Sig. (2-tailed) N |
0.129 0.538 25 |
0.035 0.869 25 |
-0.247 0.234 25 |
1
25 |
|
|
|
No of orifices |
Pearson Correlation Sig. (2-tailed) N |
0.324* 0.114 25 |
-0.174 0.404 25 |
-0.089 0.672 25 |
0.491* 0.013 25 |
1
25 |
|
|
Nest weight |
Pearson Correlation Sig. (2-tailed) N |
-0.293 0.155 25 |
-0.356 0.081 25 |
0.205 0.326 25 |
0.455* 0.022 25 |
0.082 0.697 25 |
1
25 |
|
Waste materials |
Pearson Correlation Sig. (2-tailed) N |
0.192 0.359 25 |
-0.132 0.530 25 |
-0.048 0.821 25 |
0.031 0.882 25 |
0.221 0.288 25 |
-0.028 0.894 25 |
1
25 |
*. Correlation is significant at the 0.05 level (2-tailed). |
Table 4. Correlation matrix of variables Active nests S. madraspadnum collected during the study period
|
NHG |
WS |
NW |
NL |
NO |
NWt |
WM |
SW |
No L |
LW |
No S |
|
NHG |
Pearson Correlation Sig. (1-tailed) N |
1
4 |
|
|
|
|
|
|
|
|
|
|
WS |
Pearson Correlation Sig. (1-tailed) N |
-0.469 0.266 4 |
1
4 |
|
|
|
|
|
|
|
|
|
NW |
Pearson Correlation Sig. (1-tailed) N |
0.298 0.351 4 |
-0.719 0.141 4 |
1
4 |
|
|
|
|
|
|
|
|
NL |
Pearson Correlation Sig. (1-tailed) N |
0.792* 0.104 4 |
-0.026 0.487 4 |
-0.346 0.327 4 |
1
4 |
|
|
|
|
|
|
|
NO |
Pearson Correlation Sig. (1-tailed) N |
0.073 0.464 4 |
0.846* 0.077 4 |
-0.596 0.202 4 |
0.423* 0.289 4 |
1
4 |
|
|
|
|
|
|
NWt |
Pearson Correlation Sig. (1-tailed) N |
0.690* 0.155 4 |
-0.310 0.345 4 |
-0.306 0.347 4 |
0.892* 0.054 4 |
0.028 0.486 4 |
1
4 |
|
|
|
|
|
WM |
Pearson Correlation Sig. (1-tailed) N |
-0.601 0.199 4 |
0.987** 0.007 4 |
-0.734 0.133 4 |
-0.143 0.429 4 |
0.748* 0.126 4 |
-0.370 0.315 4 |
1
4 |
|
|
|
|
SW |
Pearson Correlation Sig. (1-tailed) N |
0.932* 0.034 4 |
-0.566 0.217 4 |
0.143 0.428 4 |
0.836* 0.082 4 |
-0.095 0.452 4 |
0.881* 0.059 4 |
-0.662 0.169 4 |
1
4 |
|
|
|
No L |
Pearson Correlation Sig. (1-tailed) N |
0.914* 0.043 4 |
-0.170 0.415 4 |
-0.114 0.443 4 |
0.970* 0.015 4 |
0.344* 0.328 4 |
0.838* 0.081 4 |
-0.303 0.349 4 |
0.902* 0.049 4 |
1
4 |
|
|
LW |
Pearson Correlation Sig. (1-tailed) N |
0.934* 0.033 4 |
-0.595 0.202 4 |
0.180 0.410 4 |
0.815* 0.092 4 |
-0.126 0.437 4 |
0.866* 0.067 4 |
-0.690 0.155 4 |
0.999** 0.000 4 |
0.888** 0.056 4 |
1
4 |
|
No S |
Pearson Correlation Sig. (1-tailed) N |
0.258 0.371 4 |
-0.914 0.043 4 |
0.913* 0.043 4 |
-0.314 0.343 4 |
-0.857 0.072 4 |
-0.093 0.453 4 |
-0.888 0.056 4 |
0.254 0.373 4 |
-0.127 0.437 4 |
0.291 0.354 4 |
1
4 |
*. Correlation is significant at the 0.05 level (1-tailed). |
||||||||||||
**. Correlation is significant at the 0.01 level (1-tailed). |
NHG- Nest height from the ground; WS- Water source; NW- Nest width; NL- Nest length; NO- No. of orifices; NWt- Nest weight; WM- Waste materials; SW- Spider weight; No L- No of larva; LW- Larval weight; No S- No of spider
Investigating nest of any fauna will give more knowledge on natural history [14-16]. Studies indicate that Sceliphron madraspatanum is made on bright surfaces (walls, under tables, etc.) covered from water and direct sunlight. Also, Naumann [17] stated that Sceliphron laetum nests are commonly discovered in indistinctly light sites or shady, covered from water and direct sunlight. He also reported Syzygium formosum nests on walls covered from rain, but well. Callan [11] regarded that Syzygium formosum nests were also in a sheltered and luminous condition. Nests' height from the ground was less than 2 feet observed and it may be due to the availability of collection points, maintenance, and short duration for construction [3]. The present investigation studied the length of the nest was less than 5 cm and cell depth or width of the nest was less than 3 cm. The mean number of orifices and cells was 6.0 and the length range was measured from 2.25 to 1.92 cm. Similar results were observed by the previous studies in Anoplius infuscatus and Episyron sp. [18]. Camillo [7] also studied Sceliphron fistularium nest characteristics and notice the cells numbers in each nest ranging from 1 to 54, their length ranging from 20.8 to 29.7 mm, and their diameter ranging from 7.6 to 11.7 mm. Nests' shape was a maximum rectangle and a few were irregular shapes. The mud nest of Sceliphron madraspatanum is similar to that of Sceliphron assimile; all members of the genus Sceliphron build rectangular mud nests made up of cells [9]. Nest substrate orientation is important for protection from the abiotic (sunlight, wind, water, etc.,) and biotic factors (Gecko, Dragonflies, Spider, etc.,). This study found the east orientation was high followed by north and south. The nest orientation is used for flight orientation of the wasp surrounding nest to capture prey [19]. The condition of the nest also indicated that about 84% of the nest was inactive and the rest was active, and further studies such as feeding in the chambers were performed.
Inactive nests, mean height from the ground, and orifices or cells were lesser than the inactive nests. Likewise, the nest length and depth of the cells or width, weight were higher than the inactive nests. A total of 24 larvae was isolated from the 20 cells and weighed around 3.15 g. Similarly, the feed also isolated and tally as 153 paralyzed individuals which belonged to order Araneae and weighed 17.27 g. Among the four active nests of Mud dauber, two nests alone have been noticed the prey items and family Salticidae were showed the highest frequency as well as diversity followed by Lycosidae and Araneidae. Sceliphron wasps have been the subject of many studies in the past about prey selection. Halder et al. [2] and Gonzaga and Vasconcellos Neto [20] have been studied the same species and revealed that spiders collected from the mud wasp nests could be identified as Neoscona odites (Simon) (Araneidae) commonly known as orb-weaver spiders; Lycosa spp. (Lycosidae) or wolf spiders; and Marpissa spp. (Salticidae) or jumping spiders all of which were found abundantly in the vegetable ecosystem.
Most species appear to predominate over orb-web spiders (such as Araneidae) [21-23], however, terricolous spiders are selected in some cases, such as Sceliphron formosum (Smith), which predominantly over Salticidae [11]. Size appears to be a significant element in bait selection [24]. The active nest shape of the mud daubers was rectangle shape and substrate orientation of the nest shows 75% west followed by 25% east orientation the substrate of the nest was preferred as the wall for the long life of the larval as well as the wasp. Previously there is no study about diversity indices of spiders in a wasp nest so the present explained diversity indices of a spider. There was not much variation among the two nests of Mud daubers. The species richness, dominance index, Simpson diversity index, Shannan’H index, and Evenness are quite the same and individuals vary among the nests. The literature indicates that few insects greedily eat spiders to survive. To date, predatory mites, flies (Acroceridae), harvestman (Pholcus phalangioides) in the rice ecosystem [25], and spiders wasp (Pompilidae) feed on spiders, as been reported. However, the current study finalized that the mud dauber wasp, S. madraspatanum, is a spider hunter for juvenile survival.
CONCLUSION
Investigating natural history and related trophic connections were the initial inspiration for nest research and still feature in most studies. Almost every nest study donates to natural history knowledge. This trend likely reflects the general shift from primarily explanatory approaches to more comparative and hypothesis-driven research going beyond the detailed study of life history that is seen across biology and ecology. Likewise the present study revealed the natural history of nest of mud-dauber wasp. It illustrate the structure and other features of the nest.
ACKNOWLEDGMENTS: The authors are thankful to Principal and Management, A.V.C. College (Autonomous), Mannampandal for successfully carrying out the findings on the college campus and providing necessary guidelines.
CONFLICT OF INTEREST: None
FINANCIAL SUPPORT: None
ETHICS STATEMENT: None
1. Pulawski W. Tachysphex. http://research.calacademy.org/ent/catalog_sphecidae/1621 (accessed March 2013).
2. Halder J, Rai AB, Kodandaram MH, Shivalingaswamy TM, Dey D. Mud wasp, Sceliphron madraspatanum (Fabricius) (Hymenoptera: Sphecidae): A threat or nature's regulation of spider fauna in the vegetable agroecosystem? J Biol Control. 2012;26(4):373-5.
3. Chatenoud L, Polidori C, Federici M, Licciardi V, Andrietti F. Mud-Ball Construction by Sceliphron Mud-Dauber Wasps (Hymenoptera: Sphecidae): A Comparative Ethnological Study. Zool Stud. 2012;51(7):937-45.
4. Budrys E. On the origin of nest-building behavior in digger wasps (Hymenoptera, Apoidea). Nor J Entomol. 2001;48:45-9.
5. Polidori C, Federici M, Pesarini C, Andrietti F. Factors affecting spider prey selection by Sceliphron mud-dauber wasps (Hymenoptera: Sphecidae) in northern Italy. Anim Biol. 2007;57(1):11-28. doi:10.1163/157075607780002005.
6. Powell EC, Taylor LA. Specialists and generalists coexist within a population of spider-hunting mud dauber wasps. Behav Ecol. 2017;28(3):890-8. doi:10.1093/beheco/arx050.
7. Camillo E. The natural history of the mud-dauber wasp Sceliphron fistularium (Hymenoptera: Sphecidae) in southeastern Brazil. Rev Biol Trop. 2002;50:127-34.
8. Hochel N, Tautz J. Nesting behaviour of the paper wasp Polistes dominula in Central Europe – a flexible system for expanding into new areas. Ecosphere. 2015;6(12):262.
9. Bohart RM, Menke AS. Sphecid Wasps of the World. A Generic Revision. University of California Press; 1976.
10. Mitchell PS, Hunt JH. Nutrient and energy assays of larval provisions and feces in the black and yellow mud dauber, Sceliphron caementarium (Drury) (Hymenoptera: Sphecidae). J Kans Entomol Soc. 1984;57:700-4.
11. Callan E. Biological observations on the mud-dauber wasps Sceliphron formosum (F. Smith) (Hymenoptera: Sphecidae). Aust Entomol. 1988;14(6):78-82.
12. Genaro JA. Sobre la nidificación de Sceliphron caementarium y primer registro de Ttypoxylon texenxe para Puerto Rico (Hymenoptera: Sphecidae). Caribb J Sci. 1996;32:243-4.
13. Sebastian PA, Peter KV. Spiders of India. Universities Press (India) Hyderabad, India; 2009. p. 395.
14. Staab M, Pufal G, Tscharntke T, Klein AM. Trap nests for bees and wasps to analyse trophic interactions in changing environments-A systematic overview and user guide. Methods Ecol Evol. 2018;9(11):2226-39. doi:10.1111/2041-210X.13070
15. Perez Bote JL, Mora Rubio C, Martinez JL, Riano TR. Nesting ecology of Polistes gallicus (Hymenoptera: Vespidae) in South-Western Spain. Eur J Entomol. 2020;117:243-51.
16. Perveen F, Shah M. Nest Architectural Patterns by Three Wasp Species (Vespa velutina, Polistes flavus and Sceliphron formosum) with Reference to Their Behavior". Int J Insect Sci. 2020;5(1). doi:10.1177/IJIS.S10737.
17. Naumann ID. The biology of mud nesting Hymenoptera (and their associates) and Isoptera in rock shelters of the Kakadu Region, Northern Territory. Aust Nat Parks Wildlife Service Spec Pub. 1983;10:127-89.
18. Andrietti F, Casiraghi M, Martinoli A, Polidori C, Montresor C. Nesting habits of two spider wasps: Anoplius infuscatus and Episyron sp. (Hymenoptera: Pompilidae), with a review of the literature. Ann Soc Entomol Fr. (n.s.). 2008;44(1):93-111.
19. Kurczewski FE, Coville RE, Schal C. Observations on the Nesting and Prey of the Solitary Wasp, Tachysphex inconspicuous, with a Review of Nesting Behavior in the T. obscuripennis species group. J Insect Sci. 2010;10(183):1-16. doi:10.1673/031.010.14143
20. Gonzaga MO, Vasconcellos Neto J. Nesting characteristics and spider (Arachinida: Araneae) captured by Auplopus Agrgutus (Hymenoptera: Pompilidae) in an Area of Atlantic forest in Southestern Brazil. Entomol News. 2006;117(3):281-7.
21. White E. Nest-building and provisioning about sex in Sceliphron spirifex L. (Sphecidae). J Anim Ecol. 1962;31:317-29.
22. Coville RE. Spider-hunting sphecid wasps. In: W. Nentwig (Ed.), Ecophysiology of Spiders; 1987. pp. 309-18. Springer-Verlag, Berlin.
23. Polidori C, Trombino L, Fumagalli C, Andrietti F. The nest of the mud dauber wasp, Sceliphron spirifex (Hymenoptera: Sphecidae): an application of geological methods to structure and brood cells contents analysis. Ital J Zool. 2005;72(2):153-9.
24. Elgar MA, Jebb M. Nest provisioning in the mud-dauber wasp Sceliphron laetum (F. Smith): body mass and taxa specific prey selection. Behaviour. 1999;136(2):147-59.
25. Jackson RR, Brassington RJ. The biology of Pholcus phalangioides (Araneae, Pholcidae): Predatory versatility, araneophagy, and aggressive mimicry. J Zool. 2009;211(2):227-38. doi:10.1111/j.1469-7998.1987.tb01531.x.