Comparative morphology of immature stages of two sympatric Tenebrionidae species, with comments on their biology Author Wagner, Grzegorz K. Author Gosik, Rafał text Zootaxa 2016 4111 3 201 222 journal article 39058 10.11646/zootaxa.4111.3.1 1811da5f-ccb0-470f-84d7-83507a0dac52 1175-5326 255382 DAFF2F08-FF7D-499F-A6C0-2FA7FB080C73 Description of mature larva and pupa of Neomida haemorrhoidalis Mature larva. Coloration ( Figs 36 – 38 ). Head and thoracic segments dark brown, abdominal segments I – II light brown, next segments light brown or dark yellow; anal lobes yellowish; legs yellowish to brown. Tergites of thoracic segments distinctly sclerotized. Tergites and sternites of abdominal segments I – VIII more pigmented than lateral parts, so body seems to be covered by stipes. Cuticle smooth. Body shape and chaetotaxy ( Fig. 44 ). Body: length 12.0– 14.5 mm , width in the widest place (metathorax or abdominal segment I) 1.3–1.5 mm , shape distinctly elongated, slightly oblate dorso-ventrally in cross section, curved. At the first sight body seems to be completely void of setae. In fact chaetotaxy is well developed but weakly observed. Setae hair-like, slim, various in length, extremely transparent. Thorax: Prothorax almost as long as wide; meso- and metathorax each approximately 1.5 times broader than long. Prothorax with two horizontal rows of 6 – 8 medium long setae each first situated close to anterior margin, the second along posterior margin; mesothorax with 6 – 8 short setae placed anteriorly, next 8 very long and 4 – 6 short setae located posteriorly. Chaetotaxy of metathorax similar to mesothorcical. A pair of long setae placed latero-ventrally on each side of all thoracic segments. Legs ( Figs. 46, 47 ) thin, elongated: forelegs some stouter than next pairs; coxa of first pair subcylindrical, of next pairs conical, trochanters, conical, more or less elongated, femur of foreleg rather massive, of mid- and hindlegs thin, tibiotarsus elongated, all sparsely covered with setae various in length; claws very thin, dark brown, with a seta placed ventrally; less than two times shorter than tibiotarsus. Abdomen: Abdominal segments I – V almost of equal length, next abdominal segments decreasing gradually to the terminal parts of the body. Abdominal segment X reduced to four anal lobes of unequal size. The ventral lobe, some elongated, bigger than remaining ones. Anus located ventrally. Tergites of segments I–VII with two horizontal rows of setae (first located along anterior margin, second situated posteriorly). First row consists only of 6 – 8 short setae, second with 6 – 8 long and similar number of short setae. Long and short setae of second rows located by turns. Sternites of abdominal segments I–VIII with 6 long setae, distributed along posterior margin of each segments. Abdominal segment IX with 4 very long and several short setae dorsally and 10 long setae placed ventro-laterally. Abdominal segment IX with a single short, conical urogomphus, weakly sclerotized, localized dorso-medially. Each of anal lobes with a very short seta ( Fig. 45 ). Head ( Fig. 48 ), width 1.30–1.40 mm , prognathous, almost oval slightly widened posteriorly, distinctly oblate dorso-ventrally. Sutura coronalis well visible, sutura frontalis reached almost to antenna, frons triangular, relatively wide. Epicranium and frons more pigmented than postepicranial and lateral area. Setae of head sparse, very long distributed, dorso-laterally in two vertical rows on each side of head. There are also several long setae placed ventrally. Antennae ( Fig. 49 ) trimerous; antennomere I conical, wide; antennomere II cylindrical as long as wide; antennomere III very slender, elongated, with one elongated and two small sensillae apically. Down parts of antennomeres I and II more pigmented than the upper parts, so both antennomeres seem to be fixed from four rings. Clypeus ( Fig. 50 –left side) convex in lateral view, transverse, trapezoidal, anterior margin strongly hollowed. Basal part of clypeus more pigmented then distal parts. A pair of long setae located postero-lateral on each side. Labrum ( Fig. 50 –left side) almost as twice long as wide, with sinuated anterior margin. Labrum with 14 very long setae dorsally. All setae placed more or less marginally. Epipharynx ( Fig. 50 —right side) with six short marginal setae; two of them placed apically, next four laterally. Mandibles ( Figs. 51 a – b) asymmetrical, strong sclerotized, bifid, in dorsal view with five setae, placed laterally; right mandible wide, with well developed, semicircular molar part and small triangular tooth close to apical part; left mandible narrow, with short cut molar part and triangular tooth located basally. Maxilla ( Fig. 52 ): stipes bears five setae equal in length, hair-like and one short, stout seta. Palpus three-segmented, segment I almost as long as wide, with short seta, segments II and III distinctly longer than wide, cylindrical, segment II with two long setae, segment III with some sensillae apically. Lacinia densely covered by numerous setae of various length. Setae on lacinia straight or with curved apical part. Labium ( Fig. 52 ) divided into prementum, mentum and submentum. Prementum almost as long as wide with slightly rounded lateral parts, with two pairs of setae placed antero-laterally; mentum subcylindrical, tapering in apical part, with two pairs of setae equal in length, located laterally; submentum centrally slightly wider than long, with a pair of long setae distally; ligula concave with two pairs of setae of various length, thick, placed apically. Labial palpi two-segmented. Pupa. Coloration ( Figs. 39 – 41 ). Body grayish to yellowish (changing its color during preservation into brown), claws dark brown, lateral processes light yellow with brown spines. Shape, chaetotaxy and lateral processes ( Figs 53 – 55 ). Body elongated, slightly curved, oblate dorsoventrally, length: 6.4–7.3, width (measured on the level of second pair of legs): 2.4–2.6 mm , chaetotaxy weakly developed, setae hair-like, straight, yellow, various in length. Gin traps of abdominal tergites I – VII well developed ( Figs 42, 43 , 56 ); on abdominal segments II – VI consist of well-developed apical spine (absent on abdominal segment I), lateral and posterior spines; serrated anterior and posterior borders with triangular, strongly teeth sclerotized and some hair sensillae (one close to lateral spine strongly elongated, much bigger than others). Lateral processes of tergite VII triangular, anterior and posterior spines absent, anterior border smooth, posterior border with some setae. Abdominal segments I – VII decreasing gradually, segment VII semicircular, IX strongly reduced. Tergite IX with a pair of elongated urogomphi, with sclerotized apical parts. Head ( Fig. 57 ) semicircular, eyes weakly visible; head irregular covered by numerous short setae; head of female medially with a pair of small tubercles, of male with a pair of very elongated tubercles, always covered by relatively long setae, clypeus transversely wrinkled, with single setae on each side; labrum wide, trapezium-shaped without setae. Pronotum ( Fig. 61 ) trapezium-shaped transversely wrinkled, its central part irregular, covered by numerous short setae, borders bear with elongated setae; hypomeron glabrous; elytrals with some short setae distributed irregular; metathoracic wing sheaths smooth, mesonotum almost twice shorter than metanotum, mesonotum with a row of very short setae medially, metanotum with two rows of weakly visible setae medially and posteriorly. First abdominal tergum as wide as mesonotum; Abdominal tergites I – VII ( Figs 62, 63 ) with horizontal rows of weakly visible short setae, situated posteriorly, tergite VII with several long setae located along posterior margin; tergite IX ( Fig. 64 ) bears some short setae; abdominal ventrites I – VII ( Fig. 58 ) with weakly visible, very short setae, distributed posteriorly, ventrite VIII with elongated setae located postero-laterally. Gonotheca ( Figs 59, 60 ) of female divided, strongly elongated, curved to outside; gonotheca of male undivided. Urogomorpi possess several short setae. Femora ( Fig. 65 ) bear three elongated setae; remaining parts of legs without setae. FIGURES 36–43. Neomida haemorrhoidalis , immature stages: 36 – 38 mature larva: 36—lateral view, 37—dorsal view, 38—ventral view; 39 – 43 pupa (female): 39—ventral view, 40—lateral view, 41—dorsal view, 42, 43—lateral processes of abdomen (gin traps). FIGURES 44–47. Neomida haemorrhoidalis , mature larva, habitus and selected body parts: 44—habitus and chaetoaxy, dorsal view (left side), ventral view (right side), 45—abdominal segments 8 – 10th (lateral view), 46—front leg, 47—mid leg (Th n—thoracic segment, Ab n—abdominal segment, ur—urogomphi, co—coxa, tr—trochanter, fe—femur, tt—tibiotarsus, cl—claw). FIGURES 48–49. Neomida haemorrhoidalis , mature larva, head and antenna: 48—head shape and chaetotaxy: dorsal aspect (left side), ventral aspect (right side), 49—right antenna, (at—antenna, cl—clypeus, lbr—labrum, md—mandible, ma—maxilla, lbm—labial complex, atn n—antennomere). FIGURES 50–52. Neomida haemorrhoidalis , mature larva, mouth parts: 50—clypeus and labrum (left side), epipharynx (right side), 51a—right mandible, 51b—left mandible, 52—maxillo-labial complex (cl—clypeus, lbr—labrum, epx—epipharynx, mpa n—maxillary palpomerae, stp—stipes, pmt—praementum, mt—mentum, smt—submentum, lac—lacinia, cls —clypeal setae, epxs —setae of epipharynx, lbrs —labral setae, lgs —ligular setae, pmts —praemental setae, mts —mental setae, mds —mandibular setae, smts —submental setae, stps —stipal setae). FIGURES 53–56. Neomida haemorrhoidalis , pupa, habitus and part of gin trap: 53—total, ventral view, 54—total (dorsal view), 55—total (lateral view), 56—lateral structure of third abdominal segment (Th n—thoracic segment, Ab n—abdominal segment, ur—urogomphi, atb—anterior border, pb—posterior border, hs—hair sensilla, as—anterior spine, ls—lateral spine, ps—posterior spine). FIGURES 57–65. Neomida haemorrhoidalis , pupa, chaetotaxy of selected body parts and sexual dimorphism: 57—chaetotaxy of head, 58—detail of Ab8 (ventral aspect), 59—ventral part of Ab8 – 9 (female), 60—ventral part of Ab8 – 9 (male), 61—detail of Th1 – 3 (dorsal view), 62—detail od Ab1 (dorsal aspect), 63—detail of Ab7 (dorsal view), 64—detail of Ab8 – 9 (dorsal view), 65—chaetotaxy of front and mid leg (ur—urogomphi, THn—thoracic segment, Abn—abdominal segment, go—gonotheca, fes —femoral setae). Some information on the larval stages of B. reticulatus was presented in a comprehensive work by Gilarov (1964) . This included the habitus of the larva (lateral view), the shape of the antenna, urogomphi, labrum, epipharynx, labium, and maxilla, and the chaetotaxy of the dorsal part of the abdominal segment. Unfortunately, all figures provided by Gilarov (1964) differed in varying degrees from our observations. Some of these differences are presented in Table 1 . TABLE 1. Differences between the Bolitophagus reticulatus larva described by Gilarov (1964) and in this paper.

Feature	B. reticulatus larva
Gilarov (1964)	presented paper
Body shape	slightly oblate dorso-ventrally	rounded in cross section
Urogomphi	pronounced, thorn-like, distinctly curved upwards	small, slightly curved backward
Chaetotaxy of the body	visible only on dorsal part of 9th abdominal segment	well-developed
Head	distinctly oblate dorso-ventrally	oval
Second antennomeres	shortened and widened, with only 1 seta	elongated, with 4 setae.
Sensillae on apical antennomere	distinctly different in size and shape	almost the same
Labrum	with 4 setae	with 6 setae
Epipharynx setae	4, conical, distinct	2, thorn-like, rather small
Maxilla	only margin of lacinia covered by setae	whole lacinia covered by setae
Maxillary palpus	without setae	with 5 setae
Ligula	conical, rounded, almost as high as labial palpi	straight, not extruded from labial palpi.
Mentum	square	trapezium-shaped
Submentum	strongly elongated	rounded

TABLE 2. Differential diagnosis of immature stages of Bolitophagus reticulatus and Neomida haemorrhoidalis .

Feature	Species
B. reticulatus	N. haemorrhoidalis
Mature larva
body shape	rounded in cross section	oblate dorso-ventrally in cross section
head	hypognathous	prognathous
setae of head	numerous, differing in length	sparse, very elongated, placed laterally
distal antennomeres	cylindrical	conical
sensillae on apical antennomere	a pair of equal length	two small and one well-developed
anterior margin of labrum	rounded	sinuated
mandible dorso-laterally	with distinct conical process	smooth
legs	rather stout	elongated
abdominal segment IX	with a pair of urogomphi	with single urogomphus
anus	terminal	ventral
Pupa
head	without tubercles	with tubercles
lateral margins of pronotum	serrated	smooth
femurs	with five or more setae	with three setae
lateral processes	conical, divided	trapezium-shaped
female gonothecae	rather stout	strongly elongated
urogomphi	elongated	rather short

Some of the characteristic features listed by Matthews et al. (2010) for the Bolitophagini tribe, such as reduced urogomphi or their absence, were contrary to the information reported by Gilarov (1964) but found confirmation in our observations. Moreover, Matthews et al. (2010) also stated that their larvae have a very lightly sclerotized body with an asymmetrical head and mandibles, which corresponds with our description of B. reticulatus larvae.