UNCORRECTED PROOFZ 389 445 Continued

View metadata, citation and similar papers at core.ac.uk brought to you by CORE DTD 5 ARTICLE IN PRESS provided by Repositorio Institucional de la Universidad de Alicante 1 57 2 58 3 59 Micron xx (xxxx) 1–12 4 60 www.elsevier.com/locate/micron 5 61 6 62 7 63 8 Comparative morphology of early stages of two Mediterranean 64 9 65 10 Sarcophaga Meigen, 1826 (Diptera; Sarcophagidae) and 66 11 67 12 a review of the feeding habits of Palaearctic species 68 13 69 14 Salima Pe´rez-Moreno, M. Angeles Marcos-Garcıá*, Santos Rojo 70 15 71 16 Instituto de la Biodiversidad (CIBIO), Universidad de Alicante, E-03080 Alicante, Spain 72 17 Received 8 May 2005; revised 29 July 2005; accepted 30 July 2005 73 18 74 19 75 20 Abstract 76 21 77 22 The third instar larva of Sarcophaga hirticrus Pandelle´, 1896 and Sarcophaga javita (Peris, Gonza´lez-Mora and Mingo, 1998) are 78 23 described and figured for the first. The use of scanning electron microscopy (SEM) has been demonstrated as an effective tool for determining 79 differences at the specific level, and is here applied. The two species are distinguished from other Sarcophaga spp. and the principal 24 80 diagnostic character states are illustrated and discussed. Comparative information on immature stages morphology of the described 25 81 Palaearctic Sarcophaga species and its feeding habits are compiled and provided in a tabulated form. 26 82 q 2005 Elsevier Ltd. All rights reserved. 27 83 28 Keywords: Sarcophagidae; Immature stages; Larval morphology; Ultrastructure; Breeding habits 84 29 85 30 86 31 1. Introduction as being parasitic in vertebrates (mammals, turtles) 87 32 (Dahlem, 1991; Valle de Sales et al., 2004). 88 33 The immature stages of the majority of dipterous families In some cases, detailed descriptions of larvae implicated 89 34 remain poorly known (Hennig, 1968). In the case of the in traumatic human myiasis (Colwell and O’Connor, 2000; 90 35 Sarcophagidae relatively few papers have appeared dedi- Zumpt, 1965) or found infesting mummified human remains 91 36 cated to the larval morphology of the family (Aspoas, 1991; have been published (Sukontason et al., 2003), but in these 92 37 Ebejer, 2000; Kirk-Spriggs, 1999, 2000, 2003; Meńdez and cases the descriptions have not been attributed to any named 93 38 Pape, 2002; Zumpt, 1965). The larval stages of many species, as adults were not obtained. Therefore, the breeding 94 39 species of Sarcophagidae are necrophagous and for this habits should be related to morphological and taxonomic 95 40 reason those species termed ‘flesh-flies’ are significant in knowledge in order to improve biological and practical 96 41 forensic entomology, being second only to the Calliphoridae considerations. 97 42 (Diptera) in terms of their usefulness. Precise knowledge Sarcophagids larvae are easily recognised at family and 98 43 and precise diagnoses of their immature stages therefore generic level, but are morphologically remarkably similar 99 44 have a very practical application in estimating the post- subgenerically and inter-specifically (Aspoas, 1991). The 100 45 mortem interval (Sukontason et al., 2003; Wells et al., use of scanning electron microscopy (SEM) to observe and 101 46 2001). Other species of Sarcophaginae and Miltogramminae photograph the morphology of dipterous larvae has enabled 102 47 103 are predatory on invertebrates (e.g. other insects, snails, observation of a suite of intra-specific differences (Colwell, 48 104 earthworms, scorpions, crabs, etc.) (Meńdez and Pape, 1989), and to characterise some aspects (pseudocephalon, 49 105 2002; Pape, 1987), and several species have been reported spinules, anterior and posterior spiracles, spiracular setae, 50 106 rim of spiracular atrium tubercles and sensilia) not easily 51 107 resolved with light microscopy (Aspoas, 1991). For this 52 108 * CorrespondingUNCORRECTED authorTel.: C34 965 903400; fax: C34 965 903815. reason Cantrell PROOF (1981), indicates that it is important to 53 109 ´ 54 E-mail address: [email protected] (M.A. Marcos-Garcıa). describe the number of papillae forming the anterior 110 spiracles, the shape of the peritreme of the posterior 55 0968-4328/$ - see front matter q 2005 Elsevier Ltd. All rights reserved. 111 56 doi:10.1016/j.micron.2005.07.013 spiracles and the cephalopharingeal skeleton. 112 JMIC 869—6/9/2005—02:48—-[-no entity-]-—163450—XML MODEL 5 – pp. 1–12 DTD 5 ARTICLE IN PRESS 2 S. Pe´rez-Moreno et al. / Micron xx (xxxx) 1–12 113 Pape et al. (2002) cite 73 species of the genus approximately 24 h and were preserved in glycerine until 169 114 Sarcophaga as occurring in the Iberian Peninsula, but use. Measurements were made using a graticule mounted in 170 115 detailed morphological descriptions of larvae based on SEM an eye piece of a binocular microscope. 171 116 have been published for only five of these, namely: Stereoscan micrographs were taken with SEM HITACHI 172 117 Sarcophaga africa (Wiedemann, 1824) (as S. cruentata S3000N operated at 20 kV. Specimens of immature stages 173 118 Meigen, 1826), Sarcophaga crassipalpis Macquart, 1838, of S. hirticrus and S. javita are deposited in the 174 119 Sarcophaga dux Thomson, 1869, Sarcophaga exuberans Entomological Collection of the University of Alicante 175 120 Pandelle´, 1896 and Sarcophaga tibialis Macquart, 1851. (CIBIO). 176 121 Sarcophaga hirticrus Pandelle´, 1896 and Sarcophaga The character states examined are as follows: the 177 122 javita (Peris, Gonza´lez-Mora and Mingo, 1998) occur pseudocephalon, the cephalopharyngeal skeleton, the 178 123 sympatrically in southern Spain, where they are relatively spinules, the anterior spiracles, the spiracular atrium and 179 124 abundant. Both species have been occasionally collected the posterior spiracles. 180 125 together from the terrestrial snail Otala punctata (Mu¨ller, Taxonomic nomenclature in this paper follows Pape 181 126 1777) (Pe´rez-Moreno, 2004). (1996). Larval terminology used in this paper follows 182 127 In this study, the 3rd instar larvae of S. hirticrus and S. Teskey (1981) and Courtney et al. (2000) for ventral organ. 183 128 javita are described for the first time. Numerous features of 184 129 the 3rd instar larvae of the two species are examined by use 185 130 of SEM, in order to determine the presence of morphologi- 3. Results 186 131 cal character states of potential taxonomic value. 187 132 A table is provided which compiles published infor- 3.1. Sarcophaga hirticrus Pandelle´, 1896 188 133 mation about described larvae of Sarcophaga species and 189 134 feeding habits (Table 1). 3.1.1. Overall appearance (Fig. 1(A)) 190 135 Newly moulted larvae are creamy white in colour. 191 136 Length 9.85G0.86 mm, maximum width 3.63G0.31 mm 192 137 2. Materials and methods (nZ10). 193 138 Larvae elongated, sub-cylindrical in cross-section with a 194 139 All larval stages of S. hirticrus and S. javita were flattened ventral surface, truncated posteriorly and tapering 195 140 obtained from a laboratory colony maintained over several toward the anterior extreme. 196 141 generations under constant conditions of 25 8C temperature Posterior surface of anal segment with a distinct cavity, 197 142 and 70–85% relative humidity, and a photoperiod of 15:9 h which contains the posterior spiracles. The surface of 198 143 L:D. The colony was originated from field-collected thoracic and abdominal segments with bands of spinules of 199 144 material obtained in Mutxamel 30SYH203506 (Alicante, subtriangular form. The inter-band areas are devoid of 200 145 SE Spain) (Pe´rez-Moreno, 2004), during studies of the life spinules (Fig. 1(A)). 201 146 cycle of these species. Adult flies of S. hirticrus were placed Anal segment with two postanal tubercles with apical 202 147 into rearing cages and provided with a diet of sugar, water sensillae. Surface of the postanal tubercles covered with 203 148 and pig’s liver. The liver provided the protein meal for adult spinules in the half basal part. 204 149 female as well as a medium for larviposition. Larvae of S. 205 150 hirticrus were placed on a bed of a fine sand to facilitate the 3.1.2. Pseudocephalon (Fig. 1(B) and (C)) 206 151 pupation. Some of these puparia were left for 24/48 h in The pseudocephalon has a pair of small antennae (a) and 207 152 order to allow their sclerotisation before being killed by two palps (p) (Fig. 1(C)). Antennae appearing with two 208 153 freezing. In the case of S. javita, alive snails were provided segments, the apical (dome) in conical shape. The palps are 209 154 to the females in order to stimulate the larviposition. The mammeliform, with concentric ribbons at the apical 210 155 larvae and pupae were extracted with the help of a hand extreme where are placed five sesilla, the three posterior 211 156 needle of the snail’s shell. ones sited in the same protuberance and nominated 212 157 Third instar larvae were selected for preservation. These maxillary palpus sensilla (mxpp). 213 158 are distinguishable from larvae of other calyptrate families At both sides of the mouth, between the inferior part of 214 159 in that the majority of species possess three nearly vertical, the palps and the maxilla, appear extensive bands of 215 160 parallel, posterior spiracular openings, usually not orien- elongated, slightly corrugated overlapping oral ridges (or). 216 161 tated toward the opening in the peritreme. For permanent Ventral organs (vo) present. 217 162 preservation, larvae were killed by immersion in cold water The strong maxillae (m) are smooth, widely curved and 218 163 and slowly boiled for approximately four minutes to distend retractable (Fig. 1B). Buccal cavity displays a pair of 219 164 them.

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