PHYSIOLOGICAL ECOLOGY Development of Black Soldier (Diptera: Stratiomyidae) Larvae Fed Dairy Manure

1 2 3 4 HEIDI M. MYERS, JEFFERY K. TOMBERLIN, BARRY D. LAMBERT, AND DAVID KATTES

Environ. Entomol. 37(1): 11Ð15 (2008) ABSTRACT Black soldier ßies, illucens L., are a common colonizer of wastes. However, all published development data for this species are from studies using artiÞcial diets. This study represents the Þrst examining black soldier ßy development on animal wastes. Additionally, this study examined the ability of black soldier ßy larvae to reduce dry matter and associated nutrients in manure. Black soldier ßy larvae were fed four rates of dairy manure to determine their effects on larval and adult life history traits. Feed rate affected larval and adult development. Those fed less ration daily weighed less than those fed a greater ration. Additionally, larvae provided the least amount of dairy manure took longer to develop to the prepupal stage; however, they needed less time to reach the adult stage. Adults resulting from larvae provided 27 g dairy manure/d lived 3Ð4 d less than those fed 70 g dairy manure. Percentage survivorship to the prepupal or adult stages did not differ across treatments. Larvae fed 27 g dairy manure daily reduced manure dry matter mass by 58%, whereas those fed 70 g daily reduced dry matter 33%. Black soldier ßy larvae were able to reduce available P by 61Ð70% and N by 30Ð50% across treatments. Based on results from this study, the black soldier ßy could be used to reduce wastes and associated nutrients in conÞned bovine facilities.

The black soldier ßy is considered a beneÞcial pard 1984). Humidity Ͼ60% and temperature Ͼ26ЊC in conÞned animal facilities for its tendency to colo- resulted in Ϸ80% egg hatch (Tomberlin and Sheppard nize animal waste (Sheppard et al. 1994). Colonization 2002). of poultry and swine manure can reduce house ßy, Development data available for the black soldier ßy Musca domestica L. (Diptera: Muscidae), populations are limited to studies using an artiÞcial diet (Furman 94Ð100% and accumulated manure mass Ϸ50% (Shep- et al. 1959, May 1961, Tomberlin et al. 2002). Conse- pard 1983). Additionally, black soldier ßy prepupae quently, predicting the black soldier ßy life cycle un- can be used as animal feed (Newton et al. 1977, Shep- der Þeld conditions in conÞned animal facilities is pard et al. 1994). based on extrapolations from these previous studies. Three core laboratory studies examined fundamen- The primary objective of this study was to examine the tal developmental biology of the black soldier ßy: life history traits of black soldier ßies fed four different Furman et al. (1959), May (1961), and Tomberlin et feed rates of dairy manure, a previously unexamined al. (2002). Black soldier ßy larvae fed a Chemical dietary medium. Additionally, reduction of dry matter Specialties ManufacturersÕ Association (CSMA) stan- and associated nutrients in dairy manure by black dard ßy larval diet reach prepupal stage in ϳ2wkat soldier ßy larvae was examined. 30ЊC (Furman et al. 1959). Prepupae migrate from the food source to pupate (Sheppard et al. 1994). Adults Materials and Methods emerge after 10Ð14 d at 27Ð30ЊC (Sheppard et al. 2002). Adult black soldier ßies do not require food to Acquisition of . The experiment was performed survive but have been found to live longer when pro- using a black soldier ßy colony established at the Texas vided with a source of water (Tomberlin et al. 2002). Agricultural Experiment Station (TAES), Stephen- Two-day-old adults mate and females oviposit 2 d ville, TX. Eggs used to establish the TAES colony after copulation (Tomberlin and Sheppard 2002). Fe- originated in 2003 from a colony at the Coastal Plain males tend to lay eggs in crevices near food sources, Experiment Station, University of Georgia, Tifton, GA. with eggs hatching after4dat27ЊC (Booth and Shep- Methods for collecting black soldier ßy eggs and rear- ing the larvae were adapted from Sheppard et al. (2002). Eggs were collected and placed in a 0.5-liter 1 Department of Animal Sciences, Tarleton State University, Ste- phenville, TX 76401. plastic container, covered with a paper towel held in 2 Corresponding author: Department of Entomology, Texas A&M place with a rubberband, and stored in E-30B Percival University, 1229 N. U.S. Highway 281, Stephenville, TX 76401 (e-mail: Growth Chambers (Percival, Boone, IA; 27ЊC, 60% [email protected]). RH, and a photoperiod of 16:8 [L:D] h) until larvae 3 Texas Agricultural Experiment Station, Stephenville, TX 76401. 4 Department of Agronomy, Agribusiness, Horticulture, and Range emerged after 4 d. Once larvae were observed, 30 g Management, Tarleton State University, Stephenville, TX 76401. Gainesville diet (Hogsette 1985) mixed with 51 ml 12 ENVIRONMENTAL ENTOMOLOGY Vol. 37, no. 1

Table 1. Percentage black soldier fly survival ؎ SE to the pupal and adult stages when fed four daily dairy manure regimens under a(4 ؍ controlled conditions (N

Daily feed Percent survival to prepupal stage Percent survival to adult stage rate (g) Total Female Male No Sex Total Female Male 27 77.33 Ϯ 4.55 36.70 Ϯ 3.86a 41.10 Ϯ 3.47a 22.20 Ϯ 6.63ABC,a 61.65 Ϯ 2.74 46.80 Ϯ 2.09 53.20 Ϯ 2.09 40 84.50 Ϯ 4.30 36.20 Ϯ 1.39ab 36.50 Ϯ 1.67a 27.30 Ϯ 2.87AB,b 63.82 Ϯ 4.92 49.90 Ϯ 0.82 50.10 Ϯ 0.82 54 78.67 Ϯ 7.95 42.20 Ϯ 2.53a 38.30 Ϯ 2.97a 19.50 Ϯ 3.50ABC,b 65.78 Ϯ 6.22 52.50 Ϯ 2.39 47.50 Ϯ 2.39 70 71.25 Ϯ 4.92 42.25 Ϯ 1.88a 41.93 Ϯ 0.59a 15.85 Ϯ 1.70AC,b 62.03 Ϯ 3.37 50.12 Ϯ 1.32 49.88 Ϯ 1.32

Means in a column followed by different capital letters are signiÞcantly different. Means for both sexes and no sex within a treatment with different lowercase letters are signiÞcantly different (P Ͻ 0.05, LSD; SAS Institute 1998). Values in columns or rows not followed by letters were not signiÞcantly different across treatments. Percentage data were arcsine transformed prior to analysis. a 27ЊC, 60% relative humidity, and a photoperiod of 16:8 (L:D) h. water was provided to the larvae. Larvae were fed 4 d and returned to the rearing chambers previously men- before their use in the experiment. Voucher speci- tioned, and longevity was recorded. Adults were pro- mens were placed in the collection at the vided 0.125 ml water daily using a 1.25-cm needle TAES, Stephenville, TX. inserted through the lid. Experimental Design. The experiment was repli- Dairy Waste Analysis. Manure Ͻ6-h-old was col- cated four times from April through December 2005. lected from a commercial dairy located in Stephen- One black soldier ßy generation supplied eggs across ville, TX. This local dairy is a commercial production treatments for each replicate. Three hundred 4-d-old facility where cows are fed a standard, total mixed larvae were placed in each of four containers (5.678- ration diet ad libitum. Collected manure was placed in liter; Sterilite, Townsend, MA), covered with a 50- by 3.8-liter Ziploc bags and stored frozen in a Ϫ20ЊC 35-cm piece of T-310 100% nylon tulle (Wal-Mart Kenmore Elite heavy duty commercial chest freezer Stores, Bentonville, AR), which was held in place with (Sears, Roebuck and Company, Chicago, IL) at the a rubberband. A lid with a 12- by 8-cm rectangle cut TAES, Stephenville, TX, for 1 wk before use. Each bag in its center also was used to cover each container. The of manure used for feeding was allowed to thaw for double coverage was to prevent larvae from escaping. Ϸ24 h before use. One hundred-gram samples were All containers were stored in the rearing chamber taken from each bag and returned to the freezer. previously described with the aforementioned envi- These “predigested” samples and the total “digested” ronmental parameters. Each container with larvae was manure volume from each treatment were shipped to randomly assigned one feeding regimen of (1) 27, (2) the Soil, Water, and Forage Testing Laboratory in 40, (3) 54, or (4) 70 g of dairy cow, Bos taurus L., College Station, TX, for analysis of dry matter, N, and manure daily. Preliminary studies indicated 27 and P mass. Digested manure was deÞned as the waste 70 g manure daily were the thresholds for black soldier product remaining in a container after removal of all ßy development and manure reduction. When 40% of larvae. Percentage dry matter was determined using the larvae in a treatment reached the prepupal stage, the gravimetric method (Franson 1989). Total P was feeding was terminated for that particular treatment. measured using the nitric acid digest method (Feagley However, daily observations were made until all lar- et al. 1994). Nitrogen was measured using the Dumas vae had reached the prepupal stage or died. combustion method (Sheldrick 1986). Immature Life History Traits. For each container, Statistical Analysis. Larval and pupal development 10 larvae were randomly selected each day, individ- times, Þnal larval, prepupal, and adult weights, and ually weighed on an Adventurer Pro balance (Ohaus, adult longevity were analyzed using the PROC Mixed Pine Brook, NJ), and returned to their respective (SAS Institute 1998). The least signiÞcant difference containers. The mean larval weight recorded on the (LSD) test (SAS Institute 1998) was used after a sig- date that 40% of the larvae within a treatment reached niÞcant F test (P Յ 0.05) to separate mean differences the prepupal stage deÞned the Þnal larval weight. To for each variable. All percentage data were arcsine- differentiate life stages, prepupae had a characteristic transformed before analysis. black cuticle, whereas larvae were white (May 1961). Prepupae were removed daily from all treatments Results and individually weighed using the balance previously mentioned. Prepupae were placed individually in Immature Life History Traits. Total black soldier ßy 35-ml cups, covered with a breathable lid (Bio-Serv, larval survival was Ͼ71% throughout these experi- Frenchtown, NJ), and returned to the growth cham- ments (Table 1). Less than 20% failed to reach the ber. Each lid was labeled with information used for prepupal stage for those fed between 27 and 54 g dairy identiÞcation. Cups containing prepupae were mon- manure, whereas those fed 70 g/d suffered 29% mor- itored daily for adult emergence. tality. Individuals reaching the prepupal stage but fail- Adult Life History Traits. Teneral adults were ing to emerge as adults were placed in a separate weighed individually, sex was determined, and pupa- category including no sex. Larvae fed either 54 (F ϭ tion time was recorded for each treatment. Adult ßies 10.60, df ϭ 2, P ϭ 0.0107) or 70 g (F ϭ 62.84, df ϭ 2, were returned to their respective 35-ml capped cup P Ͻ 0.0001) had a signiÞcant difference in survivorship February 2008 MYERS ET AL.: BLACK SOLDIER FLY DEVELOPMENT 13 eeaaye sn rcMxd( Mixed Proc using analyzed were al feed Daily ae(g) rate c b a to the prepupal stage across sexes including no sex. sex the because data sex combined for only determined was column, within letters capital different by indicated difference, SigniÞcant al .Autadlra lc ode ylf-itr traits life-history fly soldier black larval and Adult 2. Table N 27 n 7FÑ31.49 Ñ F 27 0FÑ28.19 Ñ F 40 4FÑ26.85 Ñ F 54 0FÑ26.43 Ñ F 70 ϭ ϭ

Percentage female survivorship to the prepupal stage Њ ,6%rltv uiiy n htpro f1: LD h. (L:D) 16:8 of photoperiod a and humidity, relative 60% C, apesize. sample was not signiÞcantly different across treatments (F ϭ replicate. 2.08, df ϭ 3, P ϭ 0.1737). Similar results were deter- mined for males (F ϭ 1.33, df ϭ 3, P ϭ 0.3245), as well 29.28 Ñ M obnd0.1428 Combined 26.83 Ñ M obnd0.1371 Combined 25.50 Ñ M obnd0.1629 Combined 25.26 Ñ M as those individuals within the no sex (F ϭ 2.27, df ϭ 0.1786 Combined ϭ 3, P 0.1497) category. Sex Treatment (F ϭ 18.90, df ϭ 3, P Ͻ 0.0001) and sex (F ϭ 10.24, df ϭ 1, P ϭ 0.0038) signiÞcantly inßuenced larval development (Table 2). Replicate was signiÞ- cant (F ϭ 15.11, df ϭ 3, P Ͻ 0.0001) in addition to a P Յ .5 S;SSIsiue1998). Institute SAS LSD; 0.05; three-way interaction between replicate, treatment, weight larval Final and sex (F ϭ 1.97, df ϭ 9, P ϭ 0.0389). Ϯ Ϯ Ϯ Sex-based data were determined for individuals suc- Ϯ (g) .06 (4 0.0056A .08 4 27.51 (4) 0.0028A .01 4 26.17 (4) 0.0051A cessfully reaching the adult stage. Male and female 25.84 (4) 0.0077B larvae reared on 54 and 70 g dairy manure required ϳ1Ð5 d less to reach the prepupal stage than those at c lower feed rates. Larvae reached the prepupal stage in ) 25Ð31 d. Final larval weight was signiÞcantly inßu-

ϭ ϭ ϭ 30.39

؎ enced by treatment (F 10.07, df 3, P 0.0031) but avldevelopment Larval not replicate (F ϭ 2.55, df ϭ 3, P ϭ 0.1206). regimens feeding manure dairy daily four fed SE Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ .18 98 0.0893 (928) 0.1128A .54(366) 0.1564 .67(4)0.0820 (346) 0.1627 .50(6)0.1107 (367) 0.1570 .58(7)0.0958 (371) 0.1568 .10 11)0.1033 (1014) 0.1110B .57(9)0.1323 (392) 0.1517 .67(6)0.1145 (363) 0.1617 .18 94 0.1234 (944) 0.1108B .51(5)0.1466 (358) 0.1581 .51(5)0.1274 (359) 0.1591 Prepupal weight was signiÞcantly different across 0.1370 (855) 0.1122B treatments (F ϭ 84.67, df ϭ 3, P Ͻ 0.0001) and sex (F ϭ (d) 52.44, df ϭ 1, P Ͻ 0.0001). Replicate was signiÞcantly different (F ϭ 11.33, df ϭ 3, P Ͻ 0.0001). An interaction between replicate, treatment, and sex was determined b (F ϭ 2.97, df ϭ 9, P ϭ 0.0016). Male and female Ϸ

prepupae fed 70 g weighed 35% more than those fed 0.0966 27 g. rpplweight Prepupal Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Pupal developmental time differed signiÞcantly Ϯ .00(6)15.87 (366) 0.0010 .07 98 15.90 (928) 0.0007A .00(4)15.93 (346) 0.0010 .00(6)16.33 (367) 0.0010 .00(7)16.19 (371) 0.0010 .07 11)16.28 (1014) 0.0007B .09(9)17.07 (392) 0.0009 .00(6)17.20 (363) 0.0010 .07 94 17.13 (944) 0.0007C .00(5)16.56 (358) 0.0010 .00(5)17.01 (359) 0.0010 across treatment (F ϭ 6.80, df ϭ 3, P ϭ 0.0018) but not 16.78 (855) 0.0007D sex (F ϭ 0.35, df ϭ 1, P ϭ 0.5607). Replicates differed (g) signiÞcantly (F ϭ 8.69, df ϭ 3, P ϭ 0.0004) within treatment. Days for pupal development increased with greater feed rates but decreased at the 70-g feed

rate. Individuals reached the adult stage between 15.8 a and 17.2 d regardless of feed rate.

Adult Life History Traits. Mean percentage failing development Pupal Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϸ Ϯ .38(6)0.0409 (366) 0.1328 .98 77 0.0372 (717) 0.0958A .31(4)0.0335 (346) 0.1381 .33(6)0.0449 (367) 0.1333 .31(7)0.0375 (371) 0.1331 .92 70 0.0412 (740) 0.0942B .28(9)0.0543 (392) 0.1288 .33(6)0.0446 (363) 0.1373 .91C(6)0.0494 (763) 0.0941BC .33(5)0.0611 (358) 0.1343 .31(5)0.0504 (359) 0.1351 to survive from larva to adult was 37% (Table 1). No 0.0558 (723) 0.0952C

signiÞcant difference was determined between per- (d) ϭ ϭ ϭ centage survival of male (F 1.44, df 3, P 0.2955) ϫ and female (F ϭ 1.44, df ϭ 3, P ϭ 0.2955) larvae to the data All variable. any for signiÞcant not was interaction rate feed adult stage across treatments, respectively. Percent- age male survival was slightly higher than female at 27 and 40 g, whereas percentage female survival was greater than male survival at the two greater feed rates. dl weight Adult Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Adult weight differed signiÞcantly across treatment Ϯ .04(6)10.21 (361) 0.0004 .03 76 9.96 (706) 0.0003A .04(4)9.72 (345) 0.0004 .04(6)11.81 (366) 0.0004 .04(6)11.90 (369) 0.0004 .03 75 11.86 (735) 0.0003A .04(8)14.96 (389) 0.0004 .04(6)14.47 (360) 0.0004 .03 70 14.71 (750) 0.0003B .04(5)12.86 (358) 0.0004 .04(5)14.42 (358) 0.0004 .03 76 13.64 (716) 0.0003C

(F ϭ 74.13, df ϭ 3, P Ͻ 0.0001), sex (F ϭ 84.53, df ϭ (g) 1, P Ͻ 0.0001), and replicate (F ϭ 6.81, df ϭ 3, P ϭ 0.0018; Table 2). Interactions between sex and treat- ment (F ϭ 9.58, df ϭ 3, P Ͻ 0.0001) and sex and replicate (F ϭ 4.15, df ϭ 3, P ϭ 0.0061) were deter- mined. As with prepupal weight, greater adult weight was recorded with increased daily feed rate (Table 2). dl longevity Adult Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Ϯ Males weighed Ϸ18% less than females within treat- Ϯ .97(360) 0.3937 .83 (703) 0.2833A .03(343) 0.4073 .90(365) 0.3930 .93(368) 0.3933 .70B(733) 0.2780AB .82(388) 0.3802 .07(360) 0.4047 .76C(749) 0.2776BC .95(358) 0.3945 .99(359) 0.3969 ment. (717) 0.2798C Daily feed rate (F ϭ 32.50, df ϭ 3, P Ͻ 0.0001) and (d) replicate (F ϭ 28.64, df ϭ 3, P Ͻ 0.0001) signiÞcantly inßuenced male and female longevity, but sex did not (F ϭ 0.21, df ϭ 1, P ϭ 0.6487). An interaction between 14 ENVIRONMENTAL ENTOMOLOGY Vol. 37, no. 1

Table 3. Mass reduction of dairy waste fed ؎ SE for four daily this study, the appropriate rate to feed black soldier ßy a larvae dairy manure was not known. Life history traits(4 ؍ regimens fed to black soldier fly larvae (N of black soldier ßies fed 54Ð70 g dairy manure/d were Pecent reduction Daily feed improved compared with larvae fed the lesser feed rate (g) Dry matter P N rate and therefore might be optimal for raising black 27 58.20 Ϯ 5.26A 70.39 Ϯ 10.04 46.40 Ϯ 13.63 soldier ßy larvae on dairy manure. 40 54.95 Ϯ 5.05A 70.90 Ϯ 9.31 50.42 Ϯ 9.04 Food quality affects insect growth rates and posi- 54 50.08 Ϯ 1.67A 67.42 Ϯ 6.91 44.30 Ϯ 5.46 70 33.18 Ϯ 3.34B 61.53 Ϯ 8.10 30.49 Ϯ 7.53 tively correlates with larval length and percentage survival (De Haas et al. 2006). Wild black soldier ßy Means in a column followed by different letters are signiÞcantly adult weights are typically greater than for specimens different (P Յ 0.05; LSD; SAS Institute 1998). Values in columns not reared in the laboratory (Tomberlin et al. 2002). These followed by capital letters were not signiÞcantly different across differences are likely because of nutritional value of treatments. Percentage data were arcsine-transformed before analy- sis. the larval resource (Tomberlin et al. 2002). Percent- a 27ЊC, 60% relative humidity, and a photoperiod of 16:8 (L:D) h. age survival to the prepupal stage was 12Ð25% less in our study than that recorded by Tomberlin et al. (2002). Larvae held in containers with their own waste sex and treatment was determined (F ϭ 2.99, df ϭ 3, might have contaminated and consequently lowered P ϭ 0.0300). the nutritional value of newly provided manure Males and females fed either 54 or 70 g lived be- (Tomberlin et al. 2002). tween 2 and 5 d longer than those on lower feed rates. Parental effects on offspring morphology, develop- Longevity of males and females in the same treatment ment, and population dynamics have been shown for were usually similar, except at 70 g, where males lived other species (Hunter and McNeil 2000). Tempera- Ϸ1.5 d longer than females. ture and photoperiod are considered to be key vari- Dairy Waste Analysis. Nutrient analysis and dry ables responsible for parental inßuence on resulting matter reduction data are presented in Table 3. Per- offspring life histories (Hunter and McNeil 2000). centage dry matter reduction was signiÞcantly differ- Accordingly, our colony was maintained in a green- ent across treatment (F ϭ 10.07, df ϭ 3, P ϭ 0.0031). house, and these same variables could be responsible Replicate had no effect (F ϭ 2.55, df ϭ 3, P ϭ 0.1206). for the signiÞcant differences across generation ob- Larvae fed 70 g dairy manure/d reduced dry matter served in our study. Many studies, however, do not Ϸ25% less than those fed 27 g. Phosphorus reduction replicate over time to encompass multiple genera- in dairy waste was not signiÞcantly different across tions, representing a signiÞcant loss of variability. Al- treatments (F ϭ 1.68, df ϭ 3, P ϭ 0.2409), but replicate though replicates in our study were reared under was signiÞcant (F ϭ 20.64, df ϭ 3, P ϭ 0.0002). Ap- identical conditions, the parents of these ßies were proximately 67% of the P present in the dry matter housed in an outdoor greenhouse where they expe- dairy manure was removed by black soldier ßy larvae rienced light cycles and temperature regimens reßec- for all treatments. N reduction was signiÞcantly dif- tive of seasonality in northcentral Texas. ferent across replicate for each treatment (F ϭ 10.11, Adults in this study lived up to 5 d longer than those df ϭ 3, P ϭ 0.0031) but not within treatment (F ϭ 2.71, raised on artiÞcial diets (Tomberlin et al. 2002). This df ϭ 3, P ϭ 0.1079). Approximately 43% reduction in increased longevity may be a result of extended larval N mass associated with dairy manure was recorded. development periods, which allowed greater time for storing nutrients. This is not the case with other in- sects. For example, adult Schistocerca americana Drury Discussion (Orthoptera: Acrididae) rely only partially on re- All prior studies examining the life history traits of serves derived during larval development (Hahn black soldier ßy larvae and adults used artiÞcial diets. 2005). In the case of H. illucens, adults are not known CSMA (Furman et al. 1959) and Gainesville diets to feed but solely depend on nutrients stored during (Hogsette 1985), which were developed for raising the larval stage (Sheppard et al. 2002). house ßies, as well as a 15% protein layer ration Black soldier ßy development is inßuenced by mois- (Tomberlin et al. 2002), have been used to raise black ture of the larval resource (Fatchurochim et al. 1989). soldier ßies. May (1961) also examined the develop- Fresh dairy manure can be as high as 87% moisture, as ment of the black soldier ßy but failed to deÞne the reported by the American Society of Agricultural En- diet composition implemented. Data presented here gineers (ASAE) (Adhikari et al. 2005). Average per- are the Þrst for black soldier ßies raised on animal centage moisture of manure in this study was Ϸ70%, manure. whereas the artiÞcial diet studies ranged from 52 to Food availability can affect larval and adult life 70% moisture (Furman et al. 1959, Tomberlin et al. history traits (Giberson and Rosenberg 1992, Hunter 2002). Fatchurochim et al. (1989) determined black and McNeil 2000). Black soldier ßy larvae are sus- soldier ßies optimally developed in manure with 40Ð pected to develop optimally on fresh manure intro- 60% moisture and percentage survival at greater mois- duced at a low rate (Sheppard 1983); however, no ture levels was signiÞcantly reduced. supporting data have been provided. Furman et al. Reduction of P and N by black soldier ßy larvae is (1959) stated a reduced food supply can extend black promising for the development of a sustainable ma- soldier ßy larval development time up to 4 mo. Before nure digestion system in conÞned animal facilities. February 2008 MYERS ET AL.: BLACK SOLDIER FLY DEVELOPMENT 15

Because the larvae of this species naturally colonize Franson, M.A.H. (ed.). 1989. Total solids dried at manure, it may prove economically more efÞcient 103Ð105ЊC: standard methods for the examination of wa- than usage of lagoons or spreading the manure as ter and wastewater. American Public Health Association, fertilizer and possibly increasing P and N levels in the Washington, DC. soil and nearby watersheds. The Texas Institute for Furman, D. P., R. D. Young, and E. P. Catts. 1959. Hermetia Applied Environmental Research (TIAER) reported illucens (Linnaeus) as a factor in the natural control of Musca domestica Linnaeus. J. Econ. Entomol. 52: 917Ð921. dairy facilities in the Erath County, TX, are respon- Giberson, D. J., and D. M. Rosenberg. 1992. Effects of tem- sible for 65% of the N and P in the upper Bosque River perature, food quantity, and nymphal rearing density on (Adhikari et al. 2005). Reducing the mass of these life-history traits of a northern population of Hexagenia nutrients at the rates found in our study could have a (Ephemeroptera: Ephemeridae). J. North Am. Benthol. great impact on water quality problems. Soc. 11: 181Ð193. Clearly delineating the expected outcome is impor- Hahn, D. A. 2005. Larval nutrition affects lipid storage and tant when devising a sustainable dairy waste manage- growth, but not protein or carbohydrate storage in newly ment plan with the black soldier ßy. If larvae produc- eclosed adults of the grasshopper Schistocerca americana. tion for sale as feed was considered the primary J. Insect Phys. 51:1210Ð1219. objective, higher manure introduction feed rates have Hogsette, J. A. 1985. New diets for production of house ßies and stable ßies (Diptera: Muscidae) in the laboratory. J. to be used acquire the most nutritious value added Econ. Entomol. 85: 2291Ð2294. product. In contrast, if greater dry matter reduction is Hunter, M. D., and J. N. McNeil. 2000. Geographic and desired, the lower manure introduction rates need to parental inßuences on diapause by a polyphagous insect be implemented. Regardless, additional research ex- . Agric. Forest Entomol. 2: 49Ð55. amining optimal development of black soldier ßy lar- May, B. M. 1961. The occurrence in New Zealand and the vae on animal manure is needed to develop an efÞcient life-history of the soldier ßy (L.) animal digestion system with this insect. (Diptera: Stratiomyidae). New Zealand J. Sci. 4: 55Ð65. Newton, G. L., C. V. Booram, R. W. Barker, and O. M. Hale. 1977. Dried Hermetia illucens larvae meal as a supple- Acknowledgments ment for swine. J. Anim. Sci. 44: 395Ð400. SAS Institute. 1998. SAS/STAT userÕs guide, version 6, 4th The authors thank W. Watson (Department of Entomol- ed. SAS Institute, Cary, NC. ogy, North Carolina State University) and J. Muir (Texas Sheldrick, B. H. 1986. 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