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ARTICLE • Pap. Avulsos Zool. 60 (spe) • 2020 • https://doi.org/10.11606/1807-0205/2020.60.special-issue.36   copy

Potential phylogenetic significance of the number of functional abdominal spiracles in beetle pupae, with focus on Staphylinoidea (Coleoptera)

Authorship SCIMAGO INSTITUTIONS RANKINGS

Abstract

The distribution of functional abdominal spiracles in pupae of Coleoptera is reviewed based on published descriptions and original observations. Aquatic Coleoptera typically have strong modifications, generally including dramatic reductions in the number of functional spiracles and often their modification into either spiracular gills or snorkels, as a response to their environment. But pupae of the great majority of Coleoptera, which are terrestrial, show broad stability across higher taxa. Most terrestrial beetles have at least the first five pairs of abdominal spiracles functional, up to and including a full set of eight pairs. However, the number is unexpectedly low in Scarabaeoidea and within Staphyliniformia, where Histeridae and all Staphylinoidea have a confirmed maximum of four pairs of spiracles. The relation between pupal size and number of functional spiracles in terrestrial pupae is explored, and it is suggested that those groups with an unexpectedly small number of functional spiracles may have passed through a “small-size bottleneck” in their ancestry. However, this hypothesis does not explain why several families of very small beetles in other groups of Coleoptera do not show a similar reduction, and little evidence was found to support a strong relation between pupal size and number of functional spiracles at lower taxonomic levels (below family). Whether pupae are exarate or obtect apparently also has little correlation with the number of functional spiracles. However, the consistency and stability of spiracular reductions in the above groups suggests that deep historical factors are involved and thus the reductions may be of phylogenetic significance. It is urged that establishing the number of functional spiracles in beetle pupae become as standard a feature of pupal descriptions as chaetotaxy and whether they are exarate or obtect.

Key-Words.
Body size; Functional spiracle; Immature; Phylogeny; Pupa

INTRODUCTION

Beetles, an enormous group of about 400,000 described species placed in more than 190 modern families, are holometabolous insects, and thus pass through four very distinct life stages: egg, larva (usually several instars), pupa and adult. Adults are the main life stage of systematic study, and are almost always the basis for the establishment of scientific names, as well as the source of a majority of systematic characters used for classification and morphology-based phylogenetic analyses, since they include a rich character set associated with movement, feeding and reproduction. Larvae, the other active and frequently encountered life stage, also have a large suite of characters that are, to a large extent, functionally independent of those of adults. This has led to the wide use of larvae by most modern systematists, especially for phylogenetic studies. The remaining two stages, eggs and pupae, are generally inactive and short-lived compared to the other two, are less often encountered and described, and so far have seldom been included in phylogenetic studies at higher taxonomic levels. For example, the most recent and comprehensive morphological phylogenetic analysis of Coleoptera as a whole (Lawrence et al., 2011Lawrence, J.F.; Ślipiński, A.; Seago, A.E.; Thayer, M.K.; Newton, A.F. & Marvaldi, A.E. 2011. Phylogeny of the Coleoptera based on morphological characters of adults and larvae. Annales Zoologici, 61(1): 1-217. DOI) used 516 phylogenetically informative characters in the analysis, but all of these were either of adults (344) or larvae (172), with no contribution from characters of eggs or pupae.

Nevertheless, eggs and pupae do possess systematically useful characters, and have been used occasionally for systematic and phylogenetic studies within smaller groups of beetles. Eggs, for example, when included in descriptions of immature stages at all, are often described in as little as two words (e.g., “ovoid, white”), but Hinton (1981Hinton, H.E. 1981. Biology of insect eggs. Oxford, Pergamon Press. v. 2., based on unpublished work of D.C.R. Lincoln) demonstrated that, within the subfamily Staphylininae of Staphylinidae, the eggs of 43 taxa could be characterized and keyed out in a way that reflected generic and higher-level relationships within this subfamily. Pupae have received much more attention, especially through the efforts of Hinton (e.g.,1946aHinton, H.E. 1946a. A new classification of insect pupae. Proceedings of the Zoological Society of London, 116(2): 282-328., bHinton, H.E. 1946b. The “gin traps” of some beetle pupae; a protective device which appears to be unknown. Transactions of the Royal Entomological Society of London, 97: 473-496. DOI, 1948Hinton, H.E. 1948. On the origin and function of the pupal stage. Transactions of the Royal Entomological Society of London, 99: 395-409. DOI, 1949Hinton, H.E. 1949. On the function, origin, and classification of pupae. Proceedings and Transactions of the South London Entomological and Natural History Society, 1947-48: 111-154., 1955aHinton, H.E. 1955a. Protective devices of endopterygote pupae. Transactions of the Society for British Entomology, 12: 49-92., 1971Hinton, H.E. 1971. Some neglected phases in metamorphosis. Proceedings of the Royal Entomological Society of London (C), 35: 55-64.) who developed a widely accepted classification of insect pupae and drew attention to a number of pupal characters of taxonomic and phylogenetic significance for beetle pupae, such as the presence and distribution of “gin traps” and other protective devices (e.g.,Hinton, 1946bHinton, H.E. 1946b. The “gin traps” of some beetle pupae; a protective device which appears to be unknown. Transactions of the Royal Entomological Society of London, 97: 473-496. DOI, 1955aHinton, H.E. 1955a. Protective devices of endopterygote pupae. Transactions of the Society for British Entomology, 12: 49-92.). According to his classification (Hinton, 1946aHinton, H.E. 1946a. A new classification of insect pupae. Proceedings of the Zoological Society of London, 116(2): 282-328., 1949Hinton, H.E. 1949. On the function, origin, and classification of pupae. Proceedings and Transactions of the South London Entomological and Natural History Society, 1947-48: 111-154.), all known Coleoptera pupae are adecticous (lack articulated mandibles that can be moved), and most are exarate (with appendages such as antennae, wings and legs free of the body, and with more or less moveable abdomen and soft cuticle), while obtect pupae (appendages molded to the body, abdomen usually immobile, and with thick cuticle; see Fig. 1) occur within several beetle groups independently. Hinton (1946aHinton, H.E. 1946a. A new classification of insect pupae. Proceedings of the Zoological Society of London, 116(2): 282-328.) also noted that most beetle pupae occur in protected cells or even specially constructed cocoons and are terrestrial, although some are aquatic or adapted to episodic flooding. Pupae are featured prominently within some broader descriptive reviews of Coleoptera immature stages, e.g., in Costa et al. (1988Costa, C.; Vanin, S.A. & Casari-Chen, S.A. 1988. Larvas de Coleoptera do Brasil. São Paulo, Museu de Zoologia, Universidade de São Paulo. 365p.), as well as in detailed comparative studies of pupae within some groups such as Cerambycidae (e.g.,Duffy, 1953Duffy, E.A.J. 1953. A monograph of the immature stages of British and imported timber beetles (Cerambycidae). London, British Museum (Natural History)., 1957Duffy, E.A.J. 1957. A monograph of the immature stages of African timber beetles (Cerambycidae). London, British Museum (Natural History)., 1960Duffy, E.A.J. 1960. A monograph of the immature stages of Neotropical timber beetles (Cerambycidae). London, British Museum (Natural History)., 1963Duffy, E.A.J. 1963. A monograph of the immature stages of Australasian timber beetles (Cerambycidae). London, British Museum (Natural History)., 1968Duffy, E.A.J. 1968. A monograph of the immature stages of Oriental timber beetles (Cerambycidae). London, British Museum (Natural History).), Chrysomelidae (e.g.,Cox, 1996Cox, M.L. 1996. The pupae of Chrysomeloidea. In: Jolivet, P. & Cox, M.L. Chrysomelidae biology. Vol. 1: The classification, phylogeny and genetics. New York, SPB Academic Publishing. p. 119-265., 1998Cox, M.L. 1998. The pupae of Chrysomeloidea and their use in phylogeny (Coleoptera). In: Biondi, M.; Daccordi, M. & Furth, D.G. Fourth International Symposium on the Chrysomelidae, 4º. Proceedings. Torino, Museo Regionale di Scienze Naturali, 1996. p. 73-90.) and Curculionoidea (e.g.,Burke, 1968Burke, H.R. 1968. Pupae of the weevil tribe Anthonomini (Coleoptera: Curculionidae). Texas A&M University, Texas Agricultural Experiment Station, Technical Monograph, 5: 1-92.; May, 1994May, B.M. 1994. An introduction to the immature stages of Australian Curculionoidea. In: Zimmerman, E.C. Australian Weevils (Coleoptera: Curculionoidea). East Melbourne, CSIRO. v. 2, pp. 365-728.). Pupal descriptions have become not only more frequent but also more detailed and standardized over time, and now routinely include such features as the system of setae, spines or other projections that support the pupa in its cell, the presence and distribution of protective devices such as gin traps, and whether the pupa is exarate or obtect. Such pupal characters have also been used in phylogenetic studies within a few taxa, e.g., in Hydradephaga (Ruhnau, 1986Ruhnau, S. 1986. Phylogenetic relations within the Hydradephaga (Coleoptera) using larval and pupal characters. Entomologica Basiliensia, 11: 231-271.), Hydrophilidae (Archangelsky, 2004bArchangelsky, M. 2004b. Higher-level phylogeny of Hydrophilinae (Coleoptera: Hydrophilidae) based on larval, pupal and adult characters. Systematic Entomology, 29(2): 188-214. DOI), Staphylinidae: Staphylinini (Staniec & Pietrykowska-Tudruj, 2019Staniec, B. & Pietrykowska-Tudruj, E. 2019. Pupae of the mega-diverse rove beetle tribe Staphylinini (Coleoptera; Staphylinidae), their traits and systematic significance. ZooKeys, 877: 133-159. DOI), Tenebrionidae (Bouchard & Steiner, 2004Bouchard, P. & Steiner, W.E., Jr. 2004. First descriptions of Coelometopini pupae (Coleoptera: Tenebrionidae) from Australia, Southeast Asia and the Pacific region, with comments on phylogenetic relationships and antipredator adaptations. Systematic Entomology, 29(1): 101-114. DOI) and Chrysomelidae (Cox, 1998Cox, M.L. 1998. The pupae of Chrysomeloidea and their use in phylogeny (Coleoptera). In: Biondi, M.; Daccordi, M. & Furth, D.G. Fourth International Symposium on the Chrysomelidae, 4º. Proceedings. Torino, Museo Regionale di Scienze Naturali, 1996. p. 73-90.).

Figure 1
Hesperus rufipennis (Gravenhorst). Pupa (A) left lateral view, (B) dorsal view, (C) detail of functional abdominal spiracle 2, (D) detail of non-functional abdominal spiracle 6. Modified from Staniec (2004bStaniec, B. 2004b. Description of the developmental stages of Hesperus rufipennis (Gravenhorst, 1802) (Coleoptera: Staphylinidae), with comments on its biology. Annales Zoologici, 54(3): 529-539.: figs. 45, 46, 49, and 50, respectively), used with permission.

The present study is not an attempt to review pupal characters in general for potential utility in systematic and phylogenetic studies, but rather is focused on one particular trait of Coleoptera pupae, the number and placement of abdominal spiracles that are functional. More than a century ago, Verhoeff (1918Verhoeff, K.W. 1918. Studien über die Organisation der Staphylinoidea, III. Zur Kenntnis der Staphyliniden-Puppen. Zeitschrift für Wissenschaftliche Insektenbiologie, 14: 42-47, 167-171.) noted that staphylinid pupae such as Philonthus decorus (Gravenhorst) and Rugilus rufipes Germar had a restricted number of functional abdominal spiracles, not the full set of eight pairs of functional spiracles that are present in larval and adult staphylinids of the same species. Verhoeff noted that the functional spiracles on abdominal segments 1-4 were evidently open and attached to functional tracheal tubes, while the more posterior spiracles were smaller and evidently closed, with vestigial tracheal tubes, or were absent (see example in Fig. 1). Paulian (1941Paulian, R. 1941. Les premiers états des Staphylinoidea (Coleoptera). Étude de morphologie comparée. Mémoires du Muséum National d’Histoire Naturelle, Nouvelle Série, 15: 1-361, pls. 1-3.), Emden (1957Emden, F.I. van. 1957. The taxonomic significance of the characters of immature insects. Annual Review of Entomology, 2: 91-106.) and Hinton (1946aHinton, H.E. 1946a. A new classification of insect pupae. Proceedings of the Zoological Society of London, 116(2): 282-328.) also noted that this phenomenon of functional spiracular reduction in beetle pupae is common, and Paulian (1941Paulian, R. 1941. Les premiers états des Staphylinoidea (Coleoptera). Étude de morphologie comparée. Mémoires du Muséum National d’Histoire Naturelle, Nouvelle Série, 15: 1-361, pls. 1-3.) and Crowson (1981Crowson, R.A. 1981. The biology of the Coleoptera. New York, Academic Press.) suggested that the distribution of this character could have some phylogenetic significance within beetles. Unfortunately, the description of the number of functional abdominal spiracles in beetle pupae has not become as standardized as some of the other pupal characteristics noted in the previous paragraph. Some authors have noted the distribution of functional spiracles within particular groups, e.g.,Duffy (1953Duffy, E.A.J. 1953. A monograph of the immature stages of British and imported timber beetles (Cerambycidae). London, British Museum (Natural History)., 1957Duffy, E.A.J. 1957. A monograph of the immature stages of African timber beetles (Cerambycidae). London, British Museum (Natural History)., 1960Duffy, E.A.J. 1960. A monograph of the immature stages of Neotropical timber beetles (Cerambycidae). London, British Museum (Natural History)., 1963Duffy, E.A.J. 1963. A monograph of the immature stages of Australasian timber beetles (Cerambycidae). London, British Museum (Natural History)., 1968Duffy, E.A.J. 1968. A monograph of the immature stages of Oriental timber beetles (Cerambycidae). London, British Museum (Natural History).) in Cerambycidae, Cox (1996Cox, M.L. 1996. The pupae of Chrysomeloidea. In: Jolivet, P. & Cox, M.L. Chrysomelidae biology. Vol. 1: The classification, phylogeny and genetics. New York, SPB Academic Publishing. p. 119-265., 1998Cox, M.L. 1998. The pupae of Chrysomeloidea and their use in phylogeny (Coleoptera). In: Biondi, M.; Daccordi, M. & Furth, D.G. Fourth International Symposium on the Chrysomelidae, 4º. Proceedings. Torino, Museo Regionale di Scienze Naturali, 1996. p. 73-90.) in Chrysomelidae and Newton & Thayer (1995Newton, A.F. & Thayer, M.K. 1995. Protopselaphinae new subfamily for Protopselaphus new genus from Malaysia, with a phylogenetic analysis and review of the Omaliine Group of Staphylinidae including Pselaphidae (Coleoptera). In: Pakaluk, J. & Ślipiński, S.A. (Eds.). Biology, phylogeny, and classification of Coleoptera: Papers celebrating the 80th birthday of Roy A. Crowson. Warszawa, Muzeum i Instytut Zoologii PAN. p. 219-320.) and Thayer (2016Thayer, M.K. 2016. Staphylinidae Latreille, 1802. In: Beutel, R.G. & Leschen, R.A.B. (Eds.). Coleoptera, Beetles, Volume 1. Morphology and Systematics (Archostemata, Adephaga, Myxophaga, Polyphaga partim). 2.ed. Berlin, Walter de Gruyter. v. 1, p. 394-442. (Handbook of Zoology: Arthropoda: Insecta).) in Staphylinidae, but it is still not routine in Coleoptera.

The purpose of this review is to provide a brief summary of the distribution of functional abdominal spiracles in beetle pupae, and explore whether there may be some higher phylogenetic relevance in this distribution. The review is focused most heavily on the beetle group Staphyliniformia, and especially the families of Staphylinoidea. This survey is based on a combination of published descriptions of pupae plus my original observations of beetle pupae that have been identified at least to family.

MATERIAL AND METHODS

It should be emphasized first that whether abdominal spiracles in pupae are functional or non-functional is not the same as whether spiracles are present or absent, since non-functional spiracles can still be conspicuous and easily confused with functional ones (see, e.g.,Cox, 1998Cox, M.L. 1998. The pupae of Chrysomeloidea and their use in phylogeny (Coleoptera). In: Biondi, M.; Daccordi, M. & Furth, D.G. Fourth International Symposium on the Chrysomelidae, 4º. Proceedings. Torino, Museo Regionale di Scienze Naturali, 1996. p. 73-90. for discussion of the difficulty). Functional spiracles have a distinct spiracular opening and, internally, have a distinct tracheal connection and (often) a spiracular closing apparatus; non-functional spiracles are generally smaller than functional ones and lack an opening, and the attached trachea is collapsed or not evident (see example in Fig. 1). Published descriptions that do not make this distinction must be interpreted with caution, because they frequently refer to the presence of spiracles rather than functionality and may include more apical non-functional spiracles in the count. Another frequent error is that spiracles on the first abdominal segment may be overlooked because they are often more or less covered by the metathoracic wings (although spiracles on the first segment may also be genuinely absent, as evidently the case in Hydrophiloidea). Finally, care must be taken to determine that the spiracular count is from the pupa proper, not from any immediately preceding phases such a pharate pupa or pre-pupa that may be immobile but still have the spiracles of the last larval instar functional, or from a pharate adult that is visible within the pupal cuticle (see, e.g.,Costa & Vanin, 1985Costa, C. & Vanin, S.A. 1985. On the concepts of “pre-pupa”, with special reference to the Coleoptera. Revista Brasileira de Zoologia, 2(6): 339-345.; Hinton, 1971Hinton, H.E. 1971. Some neglected phases in metamorphosis. Proceedings of the Royal Entomological Society of London (C), 35: 55-64.).

More than 840 publications containing descriptions and/or illustrations of beetle pupae were consulted. This literature survey is undoubtedly more complete for the families of Staphyliniformia, because I have maintained a card catalog of immature stages of this group and sought the relevant publications for several decades. Outside of this group, the survey is based largely on a literature search using general resources such as the Zoological Record and Google Scholar, although this approach is no doubt very incomplete in detecting pupal descriptions because pupae are often not mentioned in titles or abstracts. For example, an excellent series of papers titled “Larvae of Neotropical Coleoptera …” plus taxon name (see examples in References by Costa et al. or Vanin et al.) often also include good pupal descriptions, but this is not evident from the titles. Recent comprehensive reviews of Coleoptera immatures (e.g.,Lawrence, 1991Lawrence, J.F. 1991. Order Coleoptera (general discussion, family key, many family treatments). In: Stehr, F.W. (Ed.). Immature Insects. Dubuque, Kendall/Hunt Publishing Co. v. 2. p. 144-658.) or Coleoptera as a whole (e.g., the recent Coleoptera volumes of the Handbook of Zoology: Beutel & Leschen, 2016Beutel, R.G. & Leschen, R.A.B. (Eds.). Coleoptera, Beetles, Volume 1. Morphology and Systematics (Archostemata, Adephaga, Myxophaga, Polyphaga partim). 2.ed. Berlin, Walter de Gruyter. 567p. (Handbook of Zoology: Arthropoda: Insecta).; Leschen et al., 2010Leschen, R.A.B.; Beutel, R.G. & Lawrence, J.F. 2010. Coleoptera, Beetles. Vol. 2: Morphology and systematics (Elateroidea, Bostrichiformia, Cucujiformia partim). Berlin, Walter de Gruyter. (Handbook of Zoology, Arthropoda: Insecta); Leschen & Beutel, 2014Leschen, R.A.B. & Beutel, R.G. 2014. Coleoptera, Beetles. Vol. 3: Morphology and systematics (Phytophaga). Berlin, Walter de Gruyter. 675p. (Handbook of Zoology, Arthropoda: Insecta)) were also consulted for pupal descriptions or citations of literature on them. Unfortunately, only a small percentage of the consulted published descriptions refer specifically to the distribution of functional spiracles in the studied pupae, although in some additional cases where the descriptions or illustrations were sufficiently detailed it was possible to infer the presence of functional spiracles (in these cases, the distribution is indicated with “?” in Table 1).

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Table 1
Distribution of functional abdominal spiracles (FASP) in Coleoptera pupae. Taxa are arranged alphabetically by suborder, series, superfamily, family, and taxon name(s), with addition of subfamily within Staphylinoidea and tribes within Staphylinidae. All modern Coleoptera families are listed even if no data were found, to highlight missing data at this level. The column “FASP” indicates the distribution of functional abdominal spiracles in pupae by segment number using Arabic numerals; a query (?) indicates doubt or an inferred distribution based on published figures or descriptions. The “Type” column indicates pupal type as exarate (Ex) or obtect (Ob), and also if the pupae are aquatic or periaquatic (Aq). “Sources” indicates published sources that are cited in References. An asterisk (*) before a taxon name indicates it is based on original personal observation (as “*orig.” in Sources) and a double asterisk (**) indicates other unpublished sources also indicated in Sources.

Examined pupae include some of those on which publications were based, but also many pupae present in the collections of the Field Museum of Natural History (FMNH), and a few borrowed from or studied in other collections. Some of these were identified by rearing from identifiable larvae or by association with emerged adults, but others were identified only by association with larvae and/or adults found together with them in the same microhabitat, in combination with the use of adult characters that are visible in most beetle pupae (e.g., the shape of the antenna, head, eyes, and pronotum, leg and tarsal segmentation, and length of elytra are often clearly evident in pupae). Only those pupae that could be reasonably identified to family are cited in this study. In many cases of examined small lightly sclerotized pupae, where the functionality was not clear externally, the pupa was temporarily lightly cleared by warming in lactic acid and then observed with a compound microscope (Leica Dialux 22) using transmitted light. Otherwise, pupae were observed with a Leica MZ-16 dissecting microscope. Most of these pupae are stored in 70% ethanol in vials, but a few were examined as dried pinned specimens, and a few small ones were also studied as permanent microscope slides.

The functional spiracles of beetle pupae, at least of those that pupate in terrestrial habitats, generally have simple annular spiracular openings, and may or may not have an internal closing apparatus on the tracheal connection (Hinton, 1949Hinton, H.E. 1949. On the function, origin, and classification of pupae. Proceedings and Transactions of the South London Entomological and Natural History Society, 1947-48: 111-154., and the surveyed literature and examined pupae). This is in contrast to the often complex and diverse structures of spiracular openings in beetle larvae (e.g.,Lawrence, 1991Lawrence, J.F. 1991. Order Coleoptera (general discussion, family key, many family treatments). In: Stehr, F.W. (Ed.). Immature Insects. Dubuque, Kendall/Hunt Publishing Co. v. 2. p. 144-658.). However, no attempt to record details about the structure of pupal spiracles was made in the current survey, nor was any attempt made to record the functionality of the mesothoracic spiracles, which are normally difficult to see in pupae and seldom described or figured. Furthermore, information about the functional abdominal spiracles of the corresponding adults and larvae of the surveyed pupae would be invaluable, but was generally not available in the case of most published pupal descriptions or many of the studied pupae, and no attempt was made to include such data in Table 1.

The classification of Coleoptera used here (Table 1) is adapted from Bouchard et al. (2011Bouchard, P.; Bousquet, Y.; Davies, A.E.; Alonso-Zarazaga, M.A.; Lawrence, J.F.; Lyal, C.H.C.; Newton, A.F.; Reid, C.A.M.; Schmitt, M.; Ślipiński, S.A. & Smith, A.B.T. 2011. Family-group names in Coleoptera (Insecta). ZooKeys, 88(1): 1-972. DOI) as updated by Lawrence (2016Lawrence, J.F. 2016. Classification (families & subfamilies). In: Beutel, R.G. & Leschen, R.A.B. (Eds.). Coleoptera, Beetles, Volume 1. Morphology and Systematics (Archostemata, Adephaga, Myxophaga, Polyphaga partim). 2.ed. Berlin, Walter de Gruyter. v. 1, p. 13-22. (Handbook of Zoology: Arthropoda: Insecta)), but includes a few more recent changes. Any general discussion below about phylogenetic relationships of various groups of beetles is done in the context of this classification plus recent phylogenetic studies, most notably the morphological phylogeny of Coleoptera in Lawrence et al. (2011Lawrence, J.F.; Ślipiński, A.; Seago, A.E.; Thayer, M.K.; Newton, A.F. & Marvaldi, A.E. 2011. Phylogeny of the Coleoptera based on morphological characters of adults and larvae. Annales Zoologici, 61(1): 1-217. DOI) and especially the molecular analyses of McKenna et al. (2015aMcKenna, D.D.; Farrell, B.D.; Caterino, M.S.; Farnum, C.W.; Hawks, D.C.; Maddison, D.R.; Seago, A.E.; Short, A.E.Z.; Newton, A.F. & Thayer, M.K. 2015a. Phylogeny and evolution of Staphyliniformia and Scarabaeiformia: forest litter as a stepping stone for diversification of nonphytophagous beetles. Systematic Entomology, 40(1): 35-60. DOI) for Staphyliniformia and Scarabaeoidea and McKenna et al. (2015bMcKenna, D.D.; Wild, A.L.; Kanda, K.; Bellamy, C.L.; Beutel, R.G.; Caterino, M.S.; Farnum, C.W.; Hawks, D.C.; Ivie, M.A.; Jameson, M.L.; Leschen, R.A.B.; Marvaldi, A.E.; McHugh, J.V.; Newton, A.F.; Robertson, J.A.; Thayer, M.K.; Whiting, M.F.; Lawrence, J.F.; Ślipiński, A.; Maddison, D.R. & Farrell, B.D. 2015b. The beetle tree of life reveals that Coleoptera survived end-Permian mass extinction to diversify during the Cretaceous terrestrial revolution. Systematic Entomology, 40(4): 835-880. DOI, 2019McKenna, D.D.; Shin, S.; Ahrens, D.; Balke, M.; Beza-Beza, C.; Clarke, D.J.; Donath, A.; Escalona, H.E.; Friedrich, F.; Letsch, H.; Liu, S.; Maddison, D.; Mayer, C.; Misof, B.; Murin, P.J.; Niehuis, O.; Peters, R.S.; Podsiadlowski, L.; Pohl, H.; Scully, E.D.; Yan, E.V.; Zhou, X.; Ślipiński, A. & Beutel, R.G. 2019. The evolution and genomic basis of beetle diversity. Proceedings of the National Academy of Sciences of the United States of America, 116(49): 24729-24737; online supplements. DOI) for Coleoptera as a whole, which generally support the monophyly of most of the listed suborders, series and superfamilies.

RESULTS

The results of this survey are summarized in Table 1 at the family level only, except that for the focus group Staphylinoidea, where the available information is more complete, subfamilies are also listed, and in Staphylinidae also tribes of some of the larger subfamilies when the pupal character could be determined. All taxa are listed alphabetically, from suborder to cited genera and species. Out of more than 840 publications including descriptions and/or figures of beetle pupae that were consulted, only about a third included explicit information about the distribution of functional spiracles, and these publications were concentrated in the six large families Staphylinidae, Scarabaeidae, Tenebrionidae, Cerambycidae, Chrysomelidae, and Curculionidae, where the indication of this characteristic has generally become a standard part of modern pupal descriptions. Original unpublished information based on my direct examination of more than 100 different available pupae is also included (taxa indicated with an asterisk in Table 1), to help confirm or expand on published information, and a few in litteris comments from others are added (indicated with a double asterisk and the name of the source).

All 195 modern beetle families are listed in Table 1, regardless of whether it was possible to determine the distribution of functional abdominal spiracles in pupae, in order to highlight the large gaps in knowledge of this character in Coleoptera. Pupae of perhaps two-thirds of those 195 families have been discovered and described in some way, but evidence for the distribution of functional abdominal spiracles in pupae could be found for only 93 of them, or 48%, in this survey. This does include at least one member of all modern suborders, series and superfamilies, with the exception of the small group Derodontiformia (two superfamilies if Nosodendridae is placed in its own superfamily Nosodendroidea, as in McKenna et al., 2019McKenna, D.D.; Shin, S.; Ahrens, D.; Balke, M.; Beza-Beza, C.; Clarke, D.J.; Donath, A.; Escalona, H.E.; Friedrich, F.; Letsch, H.; Liu, S.; Maddison, D.; Mayer, C.; Misof, B.; Murin, P.J.; Niehuis, O.; Peters, R.S.; Podsiadlowski, L.; Pohl, H.; Scully, E.D.; Yan, E.V.; Zhou, X.; Ślipiński, A. & Beutel, R.G. 2019. The evolution and genomic basis of beetle diversity. Proceedings of the National Academy of Sciences of the United States of America, 116(49): 24729-24737; online supplements. DOI), but in some groups like the suborder Archostemata and the polyphagan superfamilies Cleroidea, Coccinelloidea and Cucujoidea the gaps at the family level are obviously large. However, this sample is enough to highlight some apparent general patterns and conclusions, which hopefully can be tested with further discoveries and with confirmation or correction of some of the questionable entries in Table 1.

As Hinton (e.g., 1946aHinton, H.E. 1946a. A new classification of insect pupae. Proceedings of the Zoological Society of London, 116(2): 282-328., 1947Hinton, H.E. 1947. On the reduction of functional spiracles in the aquatic larvae of the Holometabola, with notes on the moulting process of spiracles. Transactions of the Royal Entomological Society of London, 98: 449-473. DOI, 1966Hinton, H.E. 1966. Respiratory adaptations of the pupae of beetles of the family Psephenidae. Philosophical Transactions of the Royal Society of London (B), 251: 211-245.) has already discussed extensively, beetle pupae and their corresponding larvae that are aquatic or inhabit periaquatic habitats, or that pupate in situations where periodic inundation is likely, generally have strong reductions in the number of functional spiracles, or other profound respiratory modifications such as the development of spiracular gills (e.g., in Myxophaga, see Reichardt & Hinton, 1976Reichardt, H. & Hinton, H.E. 1976. On the New World beetles of the family Hydroscaphidae. Papéis Avulsos de Zoologia, São Paulo, 30(1): 1-24.) or snorkels (within Scirtidae, see Jorge et al., 2019Jorge, G.; Libonatti, M.L.; Benetti, C.J. & Hamada, N. 2019. Description of the mature larva and pupa of Ora semibrunnea Pic (Coleoptera: Scirtidae) with notes on its biology. Zootaxa, 4551(1): 53-66. DOI). Indeed, Hinton (1955bHinton, H.E. 1955b. On the respiratory adaptations, biology, and taxonomy of the Psephenidae, with notes on some related families (Coleoptera). Proceedings of the Zoological Society of London, 125: 543-568., 1966Hinton, H.E. 1966. Respiratory adaptations of the pupae of beetles of the family Psephenidae. Philosophical Transactions of the Royal Society of London (B), 251: 211-245.) argued that the variation in the distribution and nature of pupal spiracles and respiration in the single small aquatic family Psephenidae, or water pennies, probably exceeds the variation in the rest of Coleoptera. A thorough review focused on these modifications and specializations in non-terrestrial beetle groups would be a worthy objective on its own, but is beyond the scope of the present study.

Here the focus is more on the pattern of functional spiracles in pupae that can be broadly considered terrestrial, a habitat that characterizes the vast majority of beetle families and higher groups. These beetles pupate in situations not likely to be flooded, such as in protected shelters under bark or in soil, sometimes in specially constructed cocoons but often in simple excavated cells (or pupal chambers), but some (e.g., within Coccinellidae and Chrysomelidae) also pupate in exposed situations such as attached to vegetation. The following Discussion explores in more detail some attributes of these terrestrial pupae, particularly the distribution of obtect pupae and the number and distribution of functional abdominal spiracles across the order.

DISCUSSION

As discussed by Hinton (1946aHinton, H.E. 1946a. A new classification of insect pupae. Proceedings of the Zoological Society of London, 116(2): 282-328., 1949Hinton, H.E. 1949. On the function, origin, and classification of pupae. Proceedings and Transactions of the South London Entomological and Natural History Society, 1947-48: 111-154.), one might expect that exposed pupae would be obtect, to provide better protection against desiccation or predation, and in fact this appears to be the case for the exposed pupae of Coccinellidae and Chrysomelidae. However (see Table 1), obtect pupae also occur in some groups that pupate in protected situations but are very small, and thus possibly more subject to desiccation (e.g., Ptiliidae, Clambidae, and Corylophidae, where the average pupal size is about 1 mm). However, not all very small beetle pupae are obtect, e.g., pupae of Hydraenidae, Sphindidae and Ciidae are also in the 1-2 mm size range but are exarate. More surprisingly, all of the many known pupae of the staphylinid subfamily Staphylininae are also obtect (Table 1). Staphylinine species are not small (known pupae range from 3-18 mm or larger), nor do they pupate in exposed situations, so an explanation in this case is not obvious (Hinton, 1946aHinton, H.E. 1946a. A new classification of insect pupae. Proceedings of the Zoological Society of London, 116(2): 282-328., 1949Hinton, H.E. 1949. On the function, origin, and classification of pupae. Proceedings and Transactions of the South London Entomological and Natural History Society, 1947-48: 111-154.). All of these multiple cases of obtect pupae in Coleoptera appear to be independently derived from exarate pupae, which are presumed to represent the ancestral condition in Coleoptera (Hinton, 1949Hinton, H.E. 1949. On the function, origin, and classification of pupae. Proceedings and Transactions of the South London Entomological and Natural History Society, 1947-48: 111-154.). In the case of Staphylininae, this derivation of obtect from exarate pupae is obvious for two reasons: (1) this subfamily is nested high within the known phylogenetic tree of Staphylinidae (see, e.g.,Grebennikov & Newton, 2009Grebennikov, V.V. & Newton, A.F. 2009. Good-bye Scydmaenidae, or why the ant-like stone beetles should become megadiverse Staphylinidae sensu latissimo (Coleoptera). European Journal of Entomology, 106(2): 275-301.; McKenna et al., 2015aMcKenna, D.D.; Farrell, B.D.; Caterino, M.S.; Farnum, C.W.; Hawks, D.C.; Maddison, D.R.; Seago, A.E.; Short, A.E.Z.; Newton, A.F. & Thayer, M.K. 2015a. Phylogeny and evolution of Staphyliniformia and Scarabaeiformia: forest litter as a stepping stone for diversification of nonphytophagous beetles. Systematic Entomology, 40(1): 35-60. DOI) while all other known pupae in Staphylinidae and Staphylinoidea are exarate, including the sister subfamily Paederinae; and (2) the observations of Frania (1992Frania, H.E. 1992. Ecdysis and other aspects of metamorphosis in rove beetles with exarate or obtect pupae (Coleoptera: Staphylinidae). Canadian Journal of Zoology, 70(12): 2326-2332.) that freshly eclosed staphylinine pupae are actually exarate, then quickly transform into obtect pupae with molded appendages and thick cuticle. However, the functional significance of obtection in this subfamily remains unclear.

If the “aquatic” groups of beetles are set aside, one can recognize in Table 1 a broad general pattern in the number of functional pupal spiracles in the terrestrial beetle groups. For most of these groups (in the suborders Adephaga and Archostemata, and the polyphagan series Bostrichiformia, Cucujiformia, Elateriformia and Scirtiformia, which together include the great majority of beetle families), functional spiracles are generally present on at least the first five abdominal segments, up to and including all eight segments in a few groups. The full range of these possibilities can be found within some of the largest of these families. For example, Cerambycidae, which probably includes the highest percentage of known pupae among beetle families thanks to the efforts of Duffy (1953Duffy, E.A.J. 1953. A monograph of the immature stages of British and imported timber beetles (Cerambycidae). London, British Museum (Natural History).-1968Duffy, E.A.J. 1968. A monograph of the immature stages of Oriental timber beetles (Cerambycidae). London, British Museum (Natural History).) and many later authors, exhibits all four states (1-5, 1-6, 1-7, 1-8), although only a single species of Pyrrhidium Fairmaire (Cerambycinae) is reported to have all eight abdominal spiracles functional (Švácha & Lawrence, 2014dŠvácha, P. & Lawrence, J.F. 2014d. Cerambycidae Latreille, 1802. In: Leschen, R.A.B. & Beutel, R.G. (Eds.). Coleoptera, Beetles. Vol. 3: Morphology and systematics: Phytophaga. Berlin, Walter de Gruyter. p. 77-177. (Handbook of Zoology, Arthropoda: Insecta)). Duffy (1960Duffy, E.A.J. 1960. A monograph of the immature stages of Neotropical timber beetles (Cerambycidae). London, British Museum (Natural History).) noted that although some cerambycid subfamilies such as Spondylidinae (as Aseminae) and Lepturinae appeared to have a constant number of functional spiracles (1-7 and 1-5, respectively), the number varied within the other subfamilies; he concluded that the number had little value as a subfamily character, and found no satisfactory correlation between the number present and the pupal environment. The other polyphagan families in these series nearly all have functional spiracles in the range from 1-5 to 1-7 within the larger families, or a fixed number within this range in the smaller families, while a full set of eight spiracles is generally rare or questionable and often in need of confirmation. The series Elateriformia, and especially its superfamily Elateroidea, have consistently higher numbers compared to the other groups, as already noted by Paulian (1941Paulian, R. 1941. Les premiers états des Staphylinoidea (Coleoptera). Étude de morphologie comparée. Mémoires du Muséum National d’Histoire Naturelle, Nouvelle Série, 15: 1-361, pls. 1-3.) and Crowson (1981Crowson, R.A. 1981. The biology of the Coleoptera. New York, Academic Press.), and this often includes the full set of 1-8. In this survey, only a few apparent exceptions were found to the presence of five or more functional spiracles in this huge assemblage of beetles, in Discolomatidae and within Erotylidae, where only four pairs of functional spiracles are found or reported for some genera (Table 1).

The distribution of functional abdominal spiracles in pupae of the two remaining large groups of beetles, the series Scarabaeiformia and Staphyliniformia, stands in strong contrast to the above pattern. Rather than a minimum of five pairs (on 1-5), a great majority of species in this large assemblage has a maximum of four pairs (1-4), down to a single pair (on 1), or rarely none. There are exceptions, notably Hydrophilidae with functional spiracles on segments 2-6, Synteliidae with possibly the same arrangement, and the possible presence of five pairs (1-5) in some Passalidae and Lucanidae (Paulian 1941Paulian, R. 1941. Les premiers états des Staphylinoidea (Coleoptera). Étude de morphologie comparée. Mémoires du Muséum National d’Histoire Naturelle, Nouvelle Série, 15: 1-361, pls. 1-3., and observed on one unidentified passalid pupa). However, most if not all of the remaining families of Scarabaeoidea, all Histeridae, and all Staphylinoidea, have a confirmed maximum of four functional spiracles (1-4). There are a few anomalous reports of higher numbers that need further exploration, e.g.,Morón (1993Morón, M.Á. 1993. Observaciones comparativas sobre la morfología pupal de los Coleoptera Melolonthidae neotropicales. Giornale Italiano di Entomologia, 6: 249-255.) noted that spiracles 1-5 or 1-6 may be open in some pupae of unnamed Cetoniinae, Morón & Salvadori (2006Morón, M.Á. & Salvadori, J.R. 2006. The third-stage larva and pupa of Demodema brevitarsis (Blanchard) (Coleoptera: Scarabaeidae: Melolonthinae) from southern Brazil. Proceedings of the Entomological Society of Washington, 108(3): 511-518.) reported that the pupa of the melolonthine genus Demodema Blanchard (now a synonym of Plectris Lepeletier & Audinet-Serville) had spiracles 5-7 open (even though they were very small compared to the clearly open 1-4), and Fuhrmann et al. (2019Fuhrmann, J.; Dias, B.M.R. & Rodrigues, S.R. 2019. Population dynamics and description of larva and pupa of Cyclocephala tucumana Bréthes, 1904 in West-Central Brazil, and remarks on immatures of other Cyclocephala species (Coleoptera: Scarabaeidae: Dynastinae). Revista Brasileira de Entomologia, 63(4): 331-342. DOI) reported that a series of pupae of the scarab species Cylcocephala tucumana Bréthes included specimens with functional spiracles on either 1-4 or 1-5. Hinton (1941aHinton, H.E. 1941a. The larva and pupa of Tachinus subterraneus (Linnaeus) (Coleoptera, Staphylinidae). Proceedings of the Royal Entomological Society of London (A), 16: 93-98.) reported that a species of Tachinus Gravenhorst (Staphylinidae; Tachyporinae) had 1-3 plus 8 open, but my own study of the pupa (in poor condition) of the related genus Tachinomorphus, which seemed similar, found that the apparent 8th spiracle likely belonged to the pharate adult inside the pupal cuticle, and the pupa of another Tachinus species described by Mank (1923Mank, H.G. 1923. The biology of the Staphylinidae. Annals of the Entomological Society of America, 16(3): 220-237.) showed conspicuous spiracles only on 1-3.

Before further discussing this unusual distribution pattern of functional pupal spiracles in beetles, it is worth considering some theoretical factors potentially relevant to the question of why pupae might benefit from reducing the number of functional spiracles compared to the corresponding larval and adult stages of the same species. First, the pupa is an inactive stage, with no food or liquid intake. Thus, its oxygen requirements should be minimal, and certainly much lower than those of the active larval and adult stages. As noted by Crowson (1981Crowson, R.A. 1981. The biology of the Coleoptera. New York, Academic Press.), Keister & Buck (1964Keister, M. & Buck, J. 1964. Respiration: some exogenous and endogenous effects on rate of respiration. In: Rockstein, M. (Ed.). The physiology of Insecta. New York, Academic Press. v. 3, p. 617-658.) demonstrated experimentally that the oxygen consumption per unit weight of the few studied beetle pupae was found to be lower than even quiescent larvae and adults at the same temperatures. Further, because there is normally no liquid intake, pupae may be highly susceptible to desiccation unless they are in a stable saturated environment. These factors would favor reducing the number of functional spiracles in pupae as compared to adults and larvae. However, such a reduction could be constrained by the size of the pupa, as can be seen from some simple calculations. If, as a first approximation, we assume the pupa is a cube, then doubling the length of the pupa while retaining the cube shape would increase its surface area by the length squared, or 4 times, and increase its volume by the length cubed, or 8 times. The pupa’s oxygen requirement would presumably be proportional to its mass (or volume), but air movement into the tracheal system would be proportional to the area of the spiracular openings (which are presumably proportional to the surface area of the pupa). In other words, with increasing pupal size, the oxygen requirement may increase faster than the ability of the tracheal system to deliver it. Therefore, larger pupae should require more, or larger, functional spiracles, or both, compared to smaller pupae, in order to meet their relatively greater oxygen needs. Thus, there is much less constraint on reduction in number of functional spiracles for small pupae than for large ones. And finally, if size increases within a group that originally has a very reduced number of functional spiracles, one might expect a secondary increase in the number of functional spiracles (which theoretically can be up to the number found in adults).

With these considerations of the potential impact of pupal size in mind, we can ask if the low numbers of functional spiracles in staphylinoids, histerids and most scarabaeoid families might be the result of one or more small-size bottlenecks in the history of these clades. According to the results of McKenna et al. (2015aMcKenna, D.D.; Farrell, B.D.; Caterino, M.S.; Farnum, C.W.; Hawks, D.C.; Maddison, D.R.; Seago, A.E.; Short, A.E.Z.; Newton, A.F. & Thayer, M.K. 2015a. Phylogeny and evolution of Staphyliniformia and Scarabaeiformia: forest litter as a stepping stone for diversification of nonphytophagous beetles. Systematic Entomology, 40(1): 35-60. DOI), these three groups are each monophyletic but do not form a clade together, so the spiracular reduction to 1-4 or less is either derived independently in each of them, or the larger number in Hydrophiloidea and possibly Synteliidae (2-6) is secondarily derived from a smaller number of no more than 1-4. Modern hydrophilid pupae include a wide size range (ca. 2-30 mm) but are so far consistent in having functional spiracles on 2-6, but the small-sized hydrophiloid family Georissidae is reported to lack functional abdominal spiracles (Hansen, 2000Hansen, M. 2000. Observations on the immature stages of Georissidae (Coleoptera: Hydrophiloidea), with remarks on the evolution of the hydrophiloid egg cocoon. Invertebrate Taxonomy, 14(6): 907-916. DOI).

The impact of small size on respiratory and other functional systems in insects in general has been explored by Polilov (2016Polilov, A.A. 2016. At the size limit - effects of miniaturization in insects. Cham, Switzerland, Springer International Publishing.), who noted that a reduction in the number of functional spiracles is a common result of extreme body size reduction in any life stage. The related idea of an ancestral small-size bottleneck has been previously proposed for the evolution of Staphylinoidea by Lawrence & Newton (1982Lawrence, J.F. & Newton, A.F., Jr. 1982. Evolution and classification of beetles. Annual Review of Ecology and Systematics, 13: 261-290.), who suggested, based on adult characters such as simplified wing venation and a reduced number of Malpighian tubules, that ancestral staphylinoids were very small, and this is also consistent with the small size of many existing staphylinoids. The low number of functional pupal spiracles in this group is consistent with this interpretation, and even consistent with the distribution pattern of functional spiracles among the included families and within Staphylinidae. In this superfamily, following McKenna et al. (2015aMcKenna, D.D.; Farrell, B.D.; Caterino, M.S.; Farnum, C.W.; Hawks, D.C.; Maddison, D.R.; Seago, A.E.; Short, A.E.Z.; Newton, A.F. & Thayer, M.K. 2015a. Phylogeny and evolution of Staphyliniformia and Scarabaeiformia: forest litter as a stepping stone for diversification of nonphytophagous beetles. Systematic Entomology, 40(1): 35-60. DOI) and prior studies, the set of relationships of families with known pupae is well established as (Hydraenidae + Ptiliidae) + ((Agyrtidae + Leiodidae) + (Silphidae + Staphylinidae)). In the first monophylum (Hydraenidae + Ptiliidae) size is always small (adults less than 1 mm to ca. 2 mm) and functional pupal spiracles are found only on the first abdominal segment, a highly unusual condition that occurs in spite of other differences between these families (Hydraenidae have semi-aquatic or aquatic larvae and adults and exarate pupae, while Ptiliidae are terrestrial with obtect pupae). The next monophylum, (Agyrtidae + Leiodidae), consistently has functional abdominal pupal spiracles on 1-2 only, even though size differences are substantial (ca. 2-3 mm for most leiodids, 10 mm or more for some agyrtids). The third and by far most diverse monophylum, (Silphidae + Staphylinidae), includes very small to very large species (less than 1 mm to over 30 mm), and includes more variation in the number of functional abdominal spiracles in pupae (1 only (rare), 1-2, 1-3, 1-4). Within this vast group, the largest species, in Silphidae and Staphylininae, consistently have spiracles 1-4 functional, while many subfamilies with smaller-sized species generally have 1-2 or 1-3 functional. The number is usually consistent within a subfamily, except for Osoriinae (where it is so far consistent within tribes) and Oxytelinae, which shows the greatest variation, even within tribes and within the single genus Bledius Leach. According to Staniec (2001aStaniec, B. 2001a. Comparative morphology of the development stages of the Polish Bledius species (Coleoptera, Staphylinidae) with comments on their biology and distribution. Lublin, Wydawnictow Uniwersytetu Marii Curie-Skłodowskiej.), most Bledius species have spiracles 1-2 functional, but some have only 1 functional (possibly related to the semi-aquatic habitat in this genus, in which the immature stages develop in burrows in salt flats or other situations where the burrows may be inundated frequently).

Histeridae also include many small species, especially in the subfamily Abraeinae, but most are medium-sized to large. Scarabaeoidea are even less likely candidates for a small-size ancestor, and modern species include some of the largest beetles, e.g., in Lucanidae, Passalidae, and especially in the scarab subfamilies Dynastinae and Cetoniinae, where pupae can exceed 60 mm in length in genera like Dynastes Kirby (Morón, 1987Morón, M.Á. 1987. Los estados inmaduros de Dynastes hyllus Chevrolat (Coleoptera: Melolonthidae; Dynastinae); con observaciones sobre su biología y el crecimiento alométrico del imago. Folia Entomológica Mexicana, 72: 33-74.). And yet, even these monstrous pupae have only abdominal spiracles 1-4 functional, although the spiracular openings are relatively huge.

Perhaps the strongest argument against a small-size bottleneck in the ancestors of staphylinoids, scarabaeoids and Histeridae as an explanation for the reduced number of functional abdominal spiracles in their pupae is the fact that very small terrestrial beetles outside of this group do not show a similar reduction. For example, the families Micromalthidae (in Archostemata) and Ciidae, Clambidae, Corylophidae and Sphindidae (in four different superfamilies of Polyphaga) all have pupae in the 1-2 mm size range, but all have at least the first five abdominal spiracles functional, regardless of whether they are obtect (Clambidae, Corylophidae) or exarate. Our knowledge about the fossil history of beetles has grown dramatically in recent years, and potentially it will be possible to compare the fossil history of staphyliniforms and scarabaeoids (which is now becoming clear, at least from the mid-Jurassic on) to the same history for the other “small-beetle” groups mentioned above, in order to compare the evolution of size in these groups and look for differences that might explain the differences in spiracular reduction. At present I can only suggest that small size may not be the main, or only, factor in determining the degree of spiracular reduction in terrestrial beetle pupae, and that one or more other yet-undetected characteristics of groups like the above staphyliniforms and scarabaeoids is a factor.

In any case, it does appear that these spiracular reductions occur consistently within staphylinoids, scarabaeoids and Histeridae, with only modest variations even across the large size ranges found within the modern members of these groups. Even though the cause may be uncertain, this consistency in turn suggests a strong correlation between this character and the phylogenetic evolution of these groups. This character may thus have some value in placing certain controversial taxa. For example, the small enigmatic family Jacobsoniidae was for a long time placed in or near Derodontoidea or its taxonomic predecessor Dermestoidea (Crowson, 1955Crowson, R.A. 1955. The natural classification of the families of Coleoptera. London, Nathaniel Lloyd. 212p. [also available as 1967 reprint, Addenda et corrigenda. Entomologist’s Monthly Magazine, 103: 209-214]., Bouchard et al., 2011Bouchard, P.; Bousquet, Y.; Davies, A.E.; Alonso-Zarazaga, M.A.; Lawrence, J.F.; Lyal, C.H.C.; Newton, A.F.; Reid, C.A.M.; Schmitt, M.; Ślipiński, S.A. & Smith, A.B.T. 2011. Family-group names in Coleoptera (Insecta). ZooKeys, 88(1): 1-972. DOI), but recent phylogenetic studies (e.g.,Lawrence et al., 2011Lawrence, J.F.; Ślipiński, A.; Seago, A.E.; Thayer, M.K.; Newton, A.F. & Marvaldi, A.E. 2011. Phylogeny of the Coleoptera based on morphological characters of adults and larvae. Annales Zoologici, 61(1): 1-217. DOI) indicated the group as a probable member of Staphylinoidea, and McKenna et al. (2015bMcKenna, D.D.; Wild, A.L.; Kanda, K.; Bellamy, C.L.; Beutel, R.G.; Caterino, M.S.; Farnum, C.W.; Hawks, D.C.; Ivie, M.A.; Jameson, M.L.; Leschen, R.A.B.; Marvaldi, A.E.; McHugh, J.V.; Newton, A.F.; Robertson, J.A.; Thayer, M.K.; Whiting, M.F.; Lawrence, J.F.; Ślipiński, A.; Maddison, D.R. & Farrell, B.D. 2015b. The beetle tree of life reveals that Coleoptera survived end-Permian mass extinction to diversify during the Cretaceous terrestrial revolution. Systematic Entomology, 40(4): 835-880. DOI, 2019) resolved Jacobsoniidae as a sister group to the monophylum (Hydraenidae + Ptiliidae). Jacobsoniid pupae are apparently unknown, but this current phylogenetic placement within Staphylinoidea leads to the prediction that they will have no more than four pairs of functional spiracles (on 1-4), and possibly only one pair as in its sister clade.

CONCLUSIONS

  1. Beetle pupae have an evidently stable set of characteristics useful for study of phylogenetic relationships at various levels.

  2. Some features, such as obtection and gin traps or other protective devices, are clearly of multiple origin within Coleoptera but probably useful phylogenetically at or below the family level.

  3. The number of functional abdominal spiracles in pupae is usually, but not necessarily, less than in larvae and adults of the same species, sometimes dramatically so in minute species (as in the clade of Hydraenidae and Ptiliidae).

  4. The number of functional abdominal spiracles may be related to body size of the most recent clade’s common ancestor. Low numbers characterize some, but not all, groups with very small species, especially those associated with aquatic habitats where they are subject to strong modifications to prevent drowning (e.g., in Myxophaga or Scirtidae), as well as certain non-aquatic groups that may have passed through a small-size bottleneck in their distant phylogenetic history, as has possibly independently occurred in staphylinoids, scarabaeoids and Histeridae.

  5. Variation in the number of functional abdominal spiracles is much lower within modern groups of family rank or below than might be predicted from theoretical considerations related to physical size. The number is often constant within large groups of species that vary enormously in size, as in most subfamilies of Scarabaeidae, in Hydrophilidae, and in “higher” Staphylinidae (Paederinae and Staphylininae). Also, the number is not invariably reduced in minute beetles, as illustrated by the presence of a minimum of 5 pairs of functional spiracles in species of Clambidae, Ciidae, Corylophidae, Micromalthidae and Sphindidae. This suggests both that historical factors outweigh functional factors in many cases, and there is the potential for high phylogenetic information content in this character.

  6. Clearly, efforts toward establishing the number of functional abdominal spiracles in more pupae of Coleoptera already seems worthwhile from a phylogenetic perspective, and describing this character should become a standard part of pupal descriptions.

ACKNOWLEDGMENTS

I thank several colleagues for providing pupal specimens for study, including Miguel Archangelsky, Claude Besuchet, Henry Frania, Vasily Grebennikov, Peter Kovarik, John Lawrence, Richard Leschen, Darren Pollock, Daniel Schmidt and Margaret Thayer, and also former curator Donald Anderson for loan of some pupae from the National History Museum in Washington DC. I also thank Federico Agrain, Caroline Chaboo, Hermant Ghante, Vladimir Gusarov and Andrew Short for help with certain literature, and Fang-Shuo Hu, Itsuro Kawashima, Margaret Thayer, Warren Steiner and the late Boris Mamaev for in litteris comments about certain pupae. I especially thank Margaret Thayer for comments and corrections on a draft of the manuscript and for arranging Fig. 1, and Bernard Staniec for permission to use the images in Fig. 1. Lastly, I thank Gabriel Biffi, Juares Fuhrmann and two anonymous reviewers for constructive comments and corrections to the submitted manuscript. Going forward, I would welcome any additions or corrections to the list in Table 1 from colleagues, whether published or in litteris. Finally, I am happy to dedicate this contribution to Dr. Cleide Costa on her 80th birthday, in acknowledgment of her lifelong contributions to the study of immature beetles, notably including her fine book on the immature beetles of Brazil (Costa et al., 1988Costa, C.; Vanin, S.A. & Casari-Chen, S.A. 1988. Larvas de Coleoptera do Brasil. São Paulo, Museu de Zoologia, Universidade de São Paulo. 365p.), and especially for featuring pupae prominently in her work and in the publications of her many colleagues and students.

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  • Published with the financial support of the "Programa de Apoio às Publicações Científicas Periódicas da USP"
Edited by: Sônia A. Casari / Gabriel Biffi

Publication Dates

  • Publication in this collection
    13 July 2020
  • Date of issue
    2020

History

  • Received
    12 Nov 2019
  • Accepted
    23 Jan 2020
  • Published
    04 Mar 2020
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