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Phylogenetic Analyses Provide New Insights Into Systematics of The 78 (1): 17 – 27 2020 © Senckenberg Gesellschaft für Naturforschung, 2020. Phylogenetic analyses provide new insights into systema- tics of the longhorned beetle tribe Acrocinini (Coleoptera: Cerambycidae: Lamiinae) Diego de Santana Souza *, 1, Tatiana Alejandra Sepúlveda 1, Luciane Marinoni 2 & Marcela Laura Monné 1 1 National Museum, Federal University of Rio de Janeiro, Department of Entomology, Quinta da Boa Vista, Sao Cristovão, 20940-040, Rio de Janeiro, Rio de Janeiro, Brazil; Diego de Santana Souza * [[email protected]]; Tatiana Alejandra Sepúlveda [tatiana. [email protected]]; Marcela Laura Monné [[email protected]] — 2 Federal University of Paraná, Department of Zoology, Caixa Postal 19020, Centro Politécnico, Jardim das Américas, 81531-990, Curitiba, Paraná, Brazil; Luciane Marinoni [[email protected]] — * Corresponding author Accepted on January 7, 2020. Published online at www.senckenberg.de/arthropod-systematics on May 26, 2020. Editors in charge: Sergio Pérez-González & Klaus-Dieter Klass Abstract. Acrocinini is a Neotropical tribe of Lamiinae (Cerambycidae) with a single species: Acrocinus longimanus (Linnaeus, 1758). Conspicuous autapomorphic characters in this species have led to divergent interpretations of the morphological body plan of the tribe and its affnities with species of Macropophora Thomson, 1864 and some species of Oreodera Audinet-Serville, 1835. Therefore, Acrocinini has not always been a monotypic tribe, and its taxonomic composition has changed according to the inclusive or exclusive placement of species of Macropophora and Oreodera. We carried out phylogenetic analyses, using maximum parsimony and Bayesian criteria, based on 34 morphological characters and 22 taxa in order to evaluate the taxonomic limits of Acrocinini and to infer the relationship among Acrocinus Illiger, 1806, Macropophora and Oreodera. Our results reveal a close relationship among these three genera, supported by six synapomorphies: prothorax with a suture surrounding the lateral tubercle; pronotum with linear coarse punctuation near posterior margin; protibia cylindrical; protibia with a projection near sulcus at inner face; protibia without a pair of apical spurs at inner margin; and protar- somere II longer than wide. New evidence confrms that Acrocinini includes Acrocinus, Macropophora and Oreodera, and increases the number of characters that defne the tribe. Additionally, we present new records of Macropophora accentifer (Olivier, 1795) for Brazil (Mato Grosso and Mato Grosso do Sul) and of Macropophora trochlearis (Linnaeus, 1758) for Venezuela and Northern Brazil (Amapá, Acre and Rondônia). Key words. Acrocinus, cladistic analysis, Harlequin, Macropophora, Oreodera, phylogeny, taxonomy. 1. Introduction Acrocinini Swainson, 1840 is a tribe of longhorned beet- morphological affnities of this species with Macropo­ les (Cerambycidae: Lamiinae) currently composed of a phora Thomson, 1864 and some species of Oreodera single species: Acrocinus longimanus (Linnaeus, 1758), Audinet-Serville, 1835. Although these genera are cur- one of the most peculiar and most colourful Neotropical rently allocated in Acanthoderini Thomson, 1860, several species of Cerambycidae, commonly known as Harle- classifcation schemes have included some of their spe- quin or Arlequim-da-mata (in Portuguese) (Fig. 1). Cur- cies as part of the tribe Acrocinini (e.g., SWAINSON 1840; rent classifcations recognize Acrocinini as a monotypic LEPESME 1946). tribe. However, its taxonomic composition has under- Originally proposed as a subfamily of Prionidae gone several changes throughout the history of classifca- (SWAINSON 1840), Acrocinini was defned by the follow- tion of Lamiinae, refecting diverse interpretations on the ing characteristics: body depressed, elytra with spines at ISSN 1863-7221 (print) | eISSN 1864-8312 (online) | DOI: 10.26049/ASP78-1-2020-02 17 Souza et al.: Phylogenetic analysis of Acrocinini Fig. 1. Male (left) and female (right) of Acrocinus longimanus (Linnaeus). Photo kindly provided by Peter Møllmann. Scale bar approxi- mately 5 cm. their apex, tarsomere I as long as the others and tarsomere Acanthoderini and left only A. longimanus in Acrocinini. III lobed or ‘heart-shaped’. Apart from Acrocinus Illiger, The composition of Acrocinini has been particularly un- 1806, SWAINSON (1840) mentioned other three genera as stable with the inclusion or exclusion of Macropophora belonging to the tribe: Macropus Audinet-Serville, 1835 and Oreodera species. Both Macropophora and Oreode­ (name synonymised under Macropophora by THOMSON ra, in addition to Acrocinus, are restricted to the Neotrop- 1864), Oreodera and Microplia Audinet-Serville, 1835. ical Region. Macropophora is a small genus composed A few years later, THOMSON (1864) redefned the system- of four species (M. accentifer, M. lacordairei, M. troch­ atic limits of Acrocinini to allocate only genera that have learis and M. worontzowi Lane, 1938) and its taxonomy small frons, fliform femora and very elongated forelegs, is relatively well-resolved (LEPESME 1946; NÉOUZE & TA- including two genera in the tribe: Acrocinus and Macropo­ VAKILIAN 2003). Oreodera, on the other hand, is one of the phora. Following THOMSON’s (1864) system, LACORDAIRE species-richest genera of the Neotropical Lamiinae with (1872) improved the diagnosis of the tribe, adding several 118 species (MONNÉ 2020) and has never been revised. characters not mentioned in previous diagnoses (e.g., the The main arguments that defend tribe-level separa- median coxal cavities widely opened and metasternum tion of Macropophora and Oreodera from Acrocinus are long) and, although without indicating characters, he as- based on modifcations related to the form of the scape sociated Acrocinini with the tribes Acanthoderini (through and femora (NÉOUZE & TAVAKILIAN 2003). However, in Oreodera) and Polyrhaphidini Thomson, 1860. the literature there is mention of characters that associate The frst taxonomic revision of Acrocinini was pub- the morphology of these three genera, suggesting they lished by LEPESME (1946), who delimited the tribe by share the same phylogenetic history, such as the shape of the scape and femora elongate and subcylindrical (nev- the mesosternal process, tibiae and tarsi (THOMSON 1860, er clavate), fore femur and tibia distinctly elongated in 1864; LEPESME 1946; LACORDAIRE 1872). In this sense, the males, and glabrous tarsi. Under this diagnosis, LEPESME unsolved systematic issues of Acrocinini may be associ- (1946) included six species in the tribe: A. longimanus, ated with two factors: (1) misinterpretation of characters Macropophora accentifer (Olivier, 1795), M. lacord­ and (2) total absence of phylogenetic studies to infer the airei Lepesme, 1946, M. trochlearis (Linnaeus, 1758), systematics of the tribe. Thus, considering the historical M. hoffmanni (Thomson, 1860) and M. lateralis Lacor- diffculties concerning the delimitation of Acrocinini, daire, 1872 (species name synonymised under M. hoff­ in this manuscript we carried out phylogenetic analyses manni, which is currently placed in Oreodera). The most based on morphological characters in order to defne the recent work dealing with systematics of Acrocinini was taxonomic limits of Acrocinini and to infer the relation- undertaken by NÉOUZE & TAVAKILIAN (2003). The authors ships among Acrocinus and the genera Macropophora considered that the diagnostic characters of Acrocinini and Oreodera. Additionally, taxonomic notes and new proposed by LEPESME (1946) were not present in all spe- distribution records are provided for species of Macro­ cies of the tribe. They then transferred Macropophora to pophora. 18 ARTHROPOD SYSTEMATICS & PHYLOGENY — 78 (1) 2020 2. Material and methods sus topology using 1,000 suboptimal trees up to one step longer. In the trees resulting from the parsimony analysis, only unambiguous character states are shown. 2.1. Material Bayesian inferences were performed in MrBayes v3.2.5 (HUELSENBECK & RONQUIST 2001) using two si- In order to infer the monophyly of Acrocinini and the phy- multaneous Markov Chain Monte Carlo runs, with 8 logenetic relationship among Acrocinus, Macropophora chains of 100 million generations each, sampling trees and Oreodera, 21 taxa were sampled, including the sole every 1,000th generation. In this analysis, the dataset was species of Acrocinini (A. longimanus), 18 representatives treated as a single partition and analysed under gamma- from 7 genera of Acanthoderini (including 3 species of distribution variation, considering all state frequencies Ma cropophora and 11 species of Oreodera), 1 species (change rates) set equal, all topologies with equal prob- of Acanthocinini Blanchard, 1845 (Acanthocinus aedilis abilities, and with unconstrained branch length. In tree (Linnaeus, 1758)), and 1 species of Polyrhaphidini (Poly­ resulting from Bayesian inference, Posterior Probability rha phis grandini Buquet, 1853; used to root the trees in (PP) was interpreted as statistical support values. the analyses) (see Table 1). We have also performed a Maximum Likelihood Outgroups were chosen based on the phylogenetic re- (ML) analysis in RAxML 7.2.6 (STAMATAKIS 2006), using lationships of Lamiinae provided by SOUZA et al. (2020) a bootstraping analysis based on 1,000 pseudoreplicates and from taxa considered close to Acrocinus in previous and the resulting tree was widely congruent with those taxonomic treatments, such as THOMSON (1860, 1864), of the Bayesian and parsimony analyses. All clades sta-
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