Zootaxa 4966 (2): 215–225 ISSN 1175-5326 (print edition) https://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2021 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4966.2.8 http://zoobank.org/urn:lsid:zoobank.org:pub:8262CD62-ACD7-482F-951F-6719394C13C8

Apertochrysa Tjeder, 1966, a new senior synonym of Tsukaguchi, 1995 (: : : )

LAURA BREITKREUZ1, PETER DUELLI2 & JOHN OSWALD3 1Museum für Naturkunde, Invalidenstraße 43, 10115 Berlin, Germany. �[email protected]; https://orcid.org/0000-0003-4776-5011 2Biodiversity and Conservation Biology, Swiss Federal Research Institute WSL, Zürcherstrasse 111, CH-8903 Birmensdorf ZH, Switzerland. �[email protected]; https://orcid.org/0000-0001-8862-8262 3Department of Entomology, Texas A&M University, College Station, Texas, USA. �[email protected]; https://orcid.org/0000-0001-7977-9309

Abstract

Apertochrysa Tjeder, 1966, is identified as a new senior subjective synonym of the large, subcosmopolitan, green lacewing genus currently known under the name Pseudomallada Tsukaguchi, 1995. Apertochrysa is rediagnosed and the taxonomic consequences of the new synonymy are reviewed. New combinations are created in Apertochrysa for 165 species formerly placed in Pseudomallada and for two species formerly placed in ; three species are transferred from Pseudomallada back to previously-established combinations in Apertochrysa; and 13 additional species are retained in Apertochrysa from previous placements. This brings the total number of Apertochrysa species to 183.

Key words: , synonymy, nomenclature, green lacewings, male terminalia

Introduction

The past 60 years has been a period of significant instability in the genus-level taxonomy and nomenclature of the green lacewing family Chrysopidae, particularly within its largest subfamily, Chrysopinae. The root cause of this instability can partially be traced to the relatively young phylogenetic history of the Chrysopidae, which comprises a clade of ca. 1400 described extant species. Recent phylogenetic research strongly supports the hypothesis that the Chrysopinae (ca. 1350 species) underwent an extensive episode of speciation during the Tertiary, with most of that radiation probably occurring during the last 40 million years of the Neogene and later Paleogene (Winterton et al., 2019; Garzón-Orduña et al., 2019; Vasilikopoulos et al., 2020). The rapid radiation that occurred during this period has left a legacy of strongly conserved gross external morphology in the adults. While notable differentiation has occurred in the morphology of the largely-concealed sclerites of the male terminalic complex, the emergence of pre-copulatory isolating mechanisms – substrate-borne vibratory signals (e.g., Henry, 1984; Henry et al. 2013) and probably also pheromonal signals (Duelli 2004, Winterton & Garzón-Orduña, 2015) – has contributed to the development of modes of speciation within the clade that are not highly dependent upon morphological differentiation. These combined factors have left a distinct stamp on the taxonomic history of the Chrysopinae. Using characters such as wing venation, coloration, and more rarely terminalia, over the 150+ year period from the mid-1700s through the 1920s early entomologists built the oldest green lacewing genus – Leach in Brewster, 1815 – into a mega-genus of over 600 species. By the mid-1930s, workers such as Tjeder (1936) and Killington (1937) had discovered the diversity of internal male terminalic traits in green lacewings and had begun to recognize their potential for the subdivision of Chrysopa. Subsequent regional studies, beginning in the 1960s (e.g., Tjeder, 1966; Hölzel, 1967), divided Chrysopa into species groups, subgenera, and new genera on the basis of terminalic traits. By the 1990s, e.g., in Brooks & Barnard’s (1990) revision of world chrysopid genera, male terminalic traits had assumed a dominant role in delimiting many

Accepted by D. Bowles: 24 Mar. 2021; published: 3 May 2021 215 green lacewing genera. An important consequence of the origin of many of the early terminalia-based genus-group names in Chrysopinae as parts of restricted regional faunas, was that neither the full geographic extent of these taxa, nor the relative phylogenetic relationships of their included species, were well known when the names were initially proposed. This situation has led in later years to the transfer of large blocks of species from one genus to another as the larger geographic distributions and phylogenetic affinities of these groups, and particularly their type species, have become better known. This paper addresses one prominent example of this taxonomic instability, and a series of related taxonomic errors. The genus hitherto named Pseudomallada Tsukaguchi, 1995 has previously carried the names Navasius* X.- k. Yang & C.-k. Yang, 1990 and Dichochrysa* X.-k. Yang, 1991 (both unavailable names). However, on the basis of recent phylogenetic work (particularly Mochizuki et al., 2017) we now know that the senior name for this genus is Apertochrysa. Here, we formally establish this new synonymy and address its taxonomic consequences. For clarity, unavailable names are flagged with asterisks (*) throughout this work.

Taxonomy

Chrysopidae: Chrysopinae: Chrysopini

Genus Apertochrysa Tjeder, 1966, Sensu Novum

Chrysopa (Apertochrysa) Tjeder, 1966: 480. Type species (of Apertochrysa): Chrysopa umbrosa Navás, 1914, by original designa- tion. Anisochrysa (Apertochrysa) Tjeder. Hölzel 1973: 341. Mallada (Triadochrysa) Adams, 1978: 294. Type species (of Triadochrysa): Mallada (Triadochrysa) triangularis Adams, 1978, by original designation. Syn. nov. Apertochrysa Tjeder. Tsukaguchi, 1985: 505. Triadochrysa Adams. Brooks & Barnard, 1990: 223 [junior synonym of Mallada]. Navasius* X.-k. Yang & C.-k. Yang, 1990: 327 [an unavailable name in the genus group; spelled identically with, but as an unavailable name not technically a nomenclatural homonym of, Navasius Esben-Petersen, 1936]. Type species: none (un- available genus-group names have no type species). Nominal genus based by X.-k. Yang & C.-k. Yang on the then-unavail- able nominal species Mallada eumorphus* X.-k. Yang & C.-k. Yang, 1990. Genus name unavailable under Art. 13.3 as its type species was not validly fixed in its original publication (its cited type species was then an unavailable nominal species and thus ineligible for fixation), and not having been proposed expressly as a new replacement name. Unavailability upheld by ICZN (2010). Syn. nov. Dichochrysa* C.-k. Yang, 1991: 150 [an unavailable name in the genus group]. Type species: none (unavailable genus-group names have no type species). Genus name unavailable under Art. 13.3 as its type species was not validly fixed in its original publication. Dichochrysa* was proposed expressly as a replacement name for Navasius* X.-k. Yang & C.-k. Yang, with which it is thus objectively synonymous. But, as no type species was originally fixed for Navasius* X.-k. Yang & C.-k. Yang, no type species could be originally fixed for its intended replacement name. Unavailability upheld by ICZN (2010). Aspöck et al., 2001:93 [Pseudomallada listed in synonymy]. Syn. nov. Pseudomallada Tsukaguchi, 1995: 67. Type species: Chrysopa cognatella Okamoto, 1914, by original designation. ICZN 2010 [Dichochrysa* in synonymy]. Syn. nov.

Diagnosis: Apertochrysa is the only chrysopid genus that possesses the following combination of male terminalic traits: (1) sternum VIII+IX simple, rounded posteriorly, (2) tignum usually present, rarely absent (then presumed secondarily lost), and (3) gonapsis always present, its form variable (as noted below). Most Apertochrysa species examined to date can be characterized by one of four generalized gonapsis morphologies (Fig. 1), which correspond closely to four clades of species identified in recent molecular phylogenetic work on the genus (Duelli et al. 2017, Fig. 1 [as Pseudomallada]). In addition, in most species of Apertochrysa the basalmost inner gradate crossvein does not arise from the pseudomedia (PsM). Distribution: Subcosmopolitan (Afrotropical, Australasian, Oceanian, Nearctic, Oriental, and Palearctic). Discussion Morphology: Historically, many of the species treated here in Apertochrysa sensu novum were formerly placed within or near the genus Mallada Navás, 1925 (e.g., Brooks & Barnard, 1990). However, recent molecular phylogenetic analyses (Mochuzuki et al., 2017; Garzón-Orduña et al., 2019; Winterton et al., 2019) have established that Apertochrysa and Mallada, while similar, are not closely related, and can be distinguished on the basis of the

216 · Zootaxa 4966 (2) © 2021 Magnolia Press BREITKREUZ et al. shape of the gonapsis (see particularly, Duelli et al., 2017), a sclerotized male terminalic element that lies medially in the posterior body wall above the (8th+)9th sternite. The gonapsis of Apertochrysa can assume a variety of forms (Fig. 1), but always consists of a pair of lateral wings, which are connected to a single anteriorly-directed rod (apodeme) (see Duelli et al. 2017 for detailed descriptions of four distinct gonapsis morphotypes found in Apertochrysa). Apart from Apertochrysa, there are only four other genera in the Chrysopinae that possess both a tignum and a gonapsis: Anomalochrysa McLachlan, 1883; Mallada Navás, 1925; Meleoma Fitch, 1855; and Peyerimhoffina Lacroix, 1920. All of these genera have gonapsis shapes that are distinctly different from Apertochrysa, and none of these genera are closely related to Apertochrysa. In the sister genera Anomalochrysa and Mallada the shape and size of the gonapsis also varies, but its form is roughly W-shaped, X-shaped, or Y-shaped. Meleoma has a small bilobed gonapsis, and in Peyerimhoffina the gonapsis is V-shaped. Apertochrysa is also morphologically similar to Steinmann; both possess a tignum (rarely absent in Apertochrysa), but Chrysoperla lacks a gonapsis (present in Apertochrysa). However, the homology and evolution of the chrysopid gonapsis is still unclear. Structures currently called gonapses are present in some members of all major clades of Chrysopini, but the structures are not present in all species of any of these clades. Current phylogenetic hypotheses for Chrysopidae are neither sufficiently complete nor robust enough to convincingly determine whether the chrysopid gonapsis evolved independently in multiple chrysopid lineages, or evolved only once, followed by independent losses in multiple chrysopid lineages. Interestingly, sclerotized male terminalic elements developed in the posterior membranous body wall below the gonarcus and above the (8th+)9th sternite are relatively rare outside the Chrysopinae, being found only in the Belonopterygini in the form of a pair of ventrolateral structures commonly referred to as parameres. Current phylogenetic hypotheses thus support the idea that the gonapsis is not a ground-plan feature of the Chrysopidae. But rather, that the structure has arisen one or more times as a non-basal evolutionary novelty within the family. This observation casts doubt on the idea that the gonapsis may represent a homolog (of sternal or some other origin) shared with other neuropteran families. More comprehensive studies of the evolution of the gonapsis, parameres and similar structures is needed to better understand this issue. Tignum state was also formerly a major diagnostic feature separating Pseudomallada (present) from Apertochrysa (absent), but recent molecular phylogenetic work suggests that tignum state is phylogenetically more variable than previously thought. In particular, secondary tignum loss may be more common in predominantly tignum-present lineages than earlier assumed, suggesting caution in using tignum presence/absence as the sole or primary character for justifying genus-group taxa in the Chrysopidae.

FIGURE 1. Representative gonapses of the four Apertochrysa species groups noted in Duelli et al. (2017), ventral view: A: prasina group, Apertochrysa ventralis (Curtis, 1834); B: alcestes group, Apertochrysa alcestes (Banks, 1911) (after Duelli et al., 2017); C: flavifrons group, Apertochrysa perfecta (Banks, 1895) (after Adams & Garland, 1982); D: venosa group, Apertochrysa venosella (Esben-Petersen, 1920) (after Tjeder, 1966). All Apertochrysa gonapses possess a median rod. The lateral “wings” vary in shape and position. Gonocristae are present only in the venosa group.

Apertochrysa, a new senior synonym of Pseudomallada Zootaxa 4966 (2) © 2021 Magnolia Press · 217 Taxonomy: Most of the pre-1990 species placed here in Apertochrysa sensu novum were originally described in the genera Chrysopa, Mallada, or Anisochrysa, and many passed through other generic combinations before being gathered into Navasius* by Yang & Yang (1990) and subsequently being transferred to Dichochrysa* X.- k. Yang, 1991. Between 1990 and 2010, ca. 55 additional new species were described in either Navasius* or Dichochrysa*, and ended up residing in Dichochrysa*. More recently, following the ICZNs (2010) ruling on the petition of Oswald (2008), which confirmed the unavailability of the generic names Navasius* and Dichochrysa*, the numerous existing Dichochrysa* combinations were rapidly replaced by new combinations in the synonymous genus Pseudomallada Tsukaguchi, 1995, which became the third most speciose genus in the family Chrysopidae, with ca. 170 species distributed nearly worldwide. This work addresses an important taxonomic issue that has arisen from the recent phylogenetic work of Duelli et al. (2017) and Mochizuki et al. (2017) on Chrysopinae. Mochizuki et al. clearly demonstrate that Apertochrysa umbrosa, the type species of the genus Apertochrysa, nests, phylogenetically, deep within a derived clade of Pseudomallada species, rendering Pseudomallada paraphyletic without (the type species of) Apertochrysa. Furthermore, because Apertochrysa has nomenclatural priority over Pseudomallada, the former name becomes a senior synonym of the latter, requiring the transfer of the numerous former Pseudomallada species into a greatly- expanded concept of Apertochrysa. Further work on Apertochrysa, especially regarding comparative morphology and species level phylogenies, is needed in order to identify and better define distinct clades within this large genus.

Taxonomy of Apertochrysa sensu novum

A list of all species included here in Apertochrysa sensu novum is given in Table 1. The table provides information on species name, author, year of publication, combination status (where new), taxonomic category, and species group placement. Species are assigned to one of four taxonomic categories, as follows: (A) Species that are currently contained in Apertochrysa, i.e., species that were placed in Apertochrysa prior to this work and are retained in that genus here. (B) Species currently included in Pseudomallada that are transferred to Apertochrysa as new combinations in this work. These transfers are required because the synonymization of Pseudomallada under Apertochrysa renders prior combinations in Pseudomallada invalid. (C) Species that are currently included in Pseudomallada, but which have pre-existing combinations in Apertochrysa. These species are transferred back to Apertochrysa in this work; they have new status, but their combinations are not new. (D) Species currently included in Mallada that are transferred to Apertochrysa as new combinations in this work. These transfers are based on published gonapsis illustrations, which support their removal from Mallada and inclusion in Apertochrysa. The gonapsis shapes of these two species are unambiguously of types C and D (Fig. 1) and cannot be confused with the gonapsis forms present in true Mallada species. Although it would be preferable to be able to physically re-examine all of the species transferred here into Apertochrysa sensu novum, this is not practicable because of the large number of species involved. We chose here to transfer all former Pseudomallada species en masse into Apertochrysa in order to make new, documented, Apertochrysa combinations available for use in on-going phylogenetic and taxonomic work, to facilitate the updating of on-line works widely utilized by the neuropterological community, and to avoid the protracted period of uncertainty regarding proper combinations that would ensue if a piecemeal transfer strategy was adopted. While some of the new Apertochrysa species will be found to belong to other genera (as was the case when they were previously placed in Pseudomallada), future removal decisions will require additional study of individual species. Brief notes on some species that may not belong in Apertochrysa are included in Table 1 to help guide future investigations. We also call attention here to two species that have recently been removed from Apertochrysa to other genera: Apertochrysa kichijoi (Kuwayama, 1936) – now Kuwayamachrysa kichijoi (Kuwayama, 1936), see Tsukaguchi & Tago 2018. Apertochrysa albolineatoides (Tsukaguchi, 1995) – now Cunctochrysa albolineatoides (Tsukaguchi, 1995), see Mochizuki et al. 2017:69.

218 · Zootaxa 4966 (2) © 2021 Magnolia Press BREITKREUZ et al. Table 1. List of species included in Apertochrysa sensu novum. Categories A-D are explained in the text. Unless otherwise annotated, Apertochrysa species group placements are based on the phylogenetic hypotheses of Duelli et al. (2017) and Mochizuki et al. (2017). Annotations: *, species group placement based on one or more published illustrations of gonapsis shape; ?, species not placed to group, or group placement tentative; —, species does not belong in Apertochrysa. Species Author(s), Year Status, Species Group Notes Category A. abdominalis (Brauer, 1856) comb. nov., B prasina group A. aegyptiaca (Navás, 1915) comb. nov., B ? A. afghanica (Hölzel, 1973) A ? tignum absent A. alarconi (Navás, 1915) comb. nov., B flavifrons group* A. albofrontata (C.-k. Yang et al., comb. nov., B ? 1999) A. alcestes (Banks, 1911) comb. nov., B alcestes group* A. alliumolens (Hölzel et al., 1997) comb. nov., B flavifrons group* A. allochroma (C.-k. Yang et al., comb. nov., B alcestes group*? 1999) A. alviolata (X.-k. Yang & C.-k. comb. nov., B ? primary type a female Yang, 1990) A. amseli (Hölzel, 1980) comb. nov., B venosa group* A. ancistroidea (X.-k. Yang & C.-k. comb. nov., B ? Yang, 1990) A. angustivittata (Dong et al., 2004) comb. nov., B ? A. anomala (Tillyard, 1917) A — Mallada sp.? A. arabica (Hölzel, 1995) comb. nov., B ? primary type a female A. arcuata (Dong et al., 2004) comb. nov., B ? A. ariadne (Hölzel, 1978) comb. nov., B prasina group* A. aromatica (C.-k. Yang & X.-k. comb. nov., B ? Yang, 1989) A. aspersa (Wesmael, 1841) comb. nov., B prasina group A. astur (Banks, 1937) comb. nov., B alcestes group A. atrosparsa (Tjeder, 1966) comb. nov., B flavifrons group* A. australis (New, 1980) A — correct genus uncertain

A. barkamana (C.-k. Yang et al., comb. nov., B ? 1992) A. baronissa (Navás, 1921) comb. nov., B flavifrons group A. basuto (Tjeder, 1966) comb. nov., B ? primary type a female A. benedictae (Séméria, 1976) comb. nov., B prasina group A. bibens (Hölzel et al., 1997) comb. nov., B venosa group* A. birungana (Navás, 1924) comb. nov., B ? A. brachychela (C.-k. Yang et al., comb. nov., B alcestes group* 1999) A. budongensis (Hölzel, 2001) comb. nov., B flavifrons group*? A. caffer (Tjeder, 1966) comb. nov., B ? primary type a female A. carinata (Dong et al., 2004) comb. nov., B flavifrons group* A. chailensis (Ghosh, 1977) comb. nov., B ? A. chaoi (C.-k. Yang et al., comb. nov., B ? 1999) ...Continued on the next page

Apertochrysa, a new senior synonym of Pseudomallada Zootaxa 4966 (2) © 2021 Magnolia Press · 219 Table 1. (Continued) Species Author(s), Year Status, Species Group Notes Category A. chlorella (Navás, 1914) comb. nov., B ? A. chloris (Schneider, 1851) comb. nov., B flavifrons group A. choui (C.-k. Yang & X.-k. comb. nov., B prasina group* Yang, 1989) A. clathrata (Schneider, 1845) comb. nov., B venosa group A. cognatella (Okamoto, 1914) comb. nov., B alcestes group* A. collartina (Navás, 1932) comb. nov., B ? A. congolana (Navás, 1911) comb. nov., B alcestes group A. cordata (X.-x. Wang & C.-k. comb. nov., B ? Yang, 1992) A. cyprina (Navás, 1932) comb. nov., B ? A. decaryna (Navás, 1924) comb. nov., B ? A. decolor (Navás, 1914) comb. nov., B alcestes group*? A. decolor (Navás, 1936) comb. nov., B ? an unreplaced junior secondary homonym of decolor (Navás, 1914) in both Pseudomallada and Apertochrysa A. deqenana (C.-k. Yang et al., comb. nov., B ? 1992) A. derbendica (Hölzel, 1967) comb. nov., B venosa group* A. deserta (Navás, 1912) comb. nov., B ? A. diaphana (C.-k. Yang et al., comb. nov., B alcestes group* 1999) A. dubia (Hölzel 1973) comb. nov., D flavifrons group* from Mallada A. duplicata (Navás, 1934) comb. nov., B alcestes group A. edwardsi (Banks, 1940) comb. nov., B alcestes group*?

A. epunctata (X.-k. Yang & C.-k. comb. nov., B ? Yang, 1990) A. eremita (Kimmins, 1955) A alcestes group*? tignum absent A. estriata (C.-k. Yang et al., comb. nov., B ? 1999) A. eumorpha (C.-k. Yang et al., comb. nov., B ? 1999) A. eurydera (Navás, 1910) stat. nov., C alcestes group tignum absent A. fanjingana (C.-k. Yang & X.-x. comb. nov., B ? Wang, 1988) A. flammefrontata (C.-k. Yang et al., comb. nov., B ? 1999) A. flavifrons (Brauer, 1851) comb. nov., B flavifrons group A. flavinotata (Dong et al., 2004) comb. nov., B ? A. flexuosa (X.-k. Yang & C.-k. comb. nov., B ? Yang, 1990) A. forcipata (X.-k. Yang & C.-k. comb. nov., B alcestes group* Yang, 1993) A. formosana (Matsumura, 1910) comb. nov., B alcestes group ...Continued on the next page

220 · Zootaxa 4966 (2) © 2021 Magnolia Press BREITKREUZ et al. Table 1. (Continued) Species Author(s), Year Status, Species Group Notes Category A. fortunata (McLachlan, 1882) comb. nov., B venosa group* A. fuscineura (C.-k. Yang et al., comb. nov., B ? 1992) A. genei (Rambur, 1842) comb. nov., B venosa group A. gradata (X.-k. Yang & C.-k. comb. nov., B alcestes group* Yang, 1993) A. granadensis (A. E. Pictet, 1865) comb. nov., B flavifrons group* A. gravesi (Navás, 1926) comb. nov., B ? A. gunvorae (Tjeder, 1966) comb. nov., B flavifrons group A. hadimensis (Canbulat & Kiyak, comb. nov., B prasina group* 2005) A. hainana (X.-k. Yang & C.-k. comb. nov., B alcestes group* Yang, 1990) A. hamata (Tjeder, 1966) comb. nov., B flavifrons group A. handschini (Navás, 1929) comb. nov., B flavifrons group A. healdi (Navás, 1926) comb. nov., B ? A. hespera (X.-k. Yang & C.-k. comb. nov., B alcestes group* Yang, 1990) A. heudei (Navás, 1934) comb. nov., B ? A. hospitalis (Hölzel & Ohm, comb. nov., B venosa group* 1995) A. huashanensis (C.-k. Yang & X.-k. comb. nov., B ? Yang, 1989) A. hubeiana (C.-k. Yang & X.-x. comb. nov., B ? Wang, 1990) A. iberica (Navás, 1903) comb. nov., B flavifrons group A. ifranina (Navás, 1936) comb. nov., B ? A. ignea (X.-k. Yang & C.-k. comb. nov., B alcestes group*? Yang, 1990) A. illota (Navás, 1908) comb. nov., B ? A. incongrua (Fraser, 1951) comb. nov., B ? A. incrassata (Tjeder, 1966) comb. nov., B flavifrons group* A. ingae (Tjeder, 1966) comb. nov., B venosa group* A. iniqua (Navás, 1931) comb. nov., B ? A. inopinata (Hölzel & Ohm, comb. nov., B flavifrons group? closely related to venosa group 1995) A. inornata (Navás, 1901) comb. nov., B ?

A. irrorella (Navás, 1936) comb. nov., B ? A. jiuzhaigouana (X.-k. Yang & X.-x. comb. nov., B prasina group* Wang in X.-k. Yang et al., 2005) A. joannisi (Navás, 1910) stat. nov., C ? tignum absent A. karooensis (Hölzel, 1993) comb. nov., B venosa group A. kiangsuensis (Navás, 1934) comb. nov., B alcestes group* ...Continued on the next page

Apertochrysa, a new senior synonym of Pseudomallada Zootaxa 4966 (2) © 2021 Magnolia Press · 221 Table 1. (Continued) Species Author(s), Year Status, Species Group Notes Category A. kibonotoensis (van der Weele, comb. nov., B flavifrons group 1910) A. kinnaurensis (Gosh, 1977) comb. nov., D venosa group* from Mallada A. leai (Tillyard, 1917) A — Mallada sp.? A. lii (C.-k. Yang et al., comb. nov., B prasina group* 1999) A. longwangshana (X.-k. Yang, 1998) comb. nov., B alcestes group*

A. lophophora (X.-k. Yang & C.-k. comb. nov., B alcestes group* Yang, 1990) A. luaboensis (Tjeder, 1966) comb. nov., B flavifrons group A. luctuosa (Banks, 1911) comb. nov., B flavifrons group A. macleodi (Adams & Garland, comb. nov., B flavifrons group 1983) A. maghrebina (Hölzel & Ohm, comb. nov., B flavifrons group* 1984) A. makrana (Hölzel, 1966) comb. nov., B venosa group* A. mangkangensis (Dong et al., 2004) comb. nov., B alcestes group*?

A. marchionissa (Navás, 1915) comb. nov., B ? A. mauriciana (Hölzel & Ohm, comb. nov., B venosa group* 1991) A. mediata (X.-k. Yang & C.-k. comb. nov., B ? Yang, 1993) A. medogana (C.-k. Yang et al. in comb. nov., B ? Huang et al., 1988) A. melanopis (Navás, 1914) comb. nov., B ? A. militaris (Hölzel & Ohm, comb. nov., B flavifrons group* 2000) A. mira (Hölzel, 1973) comb. nov., B flavifrons group* A. namibensis (Hölzel, 1993) comb. nov., B venosa group* A. nautarum (Tillyard, 1917) A — Mallada sp.? A. nicolaina (Navás, 1929) comb. nov., B venosa group A. nigra (McLachlan, 1869) comb. nov., B ? A. nigricornuta (X.-k. Yang & C.-k. comb. nov., B ? Yang, 1990) A. norfolkensis (Tillyard, 1917) A — Mallada sp.? A. nubilata (Navás, 1910) comb. nov., B alcestes group A. nyassalandica (Navás, 1914) comb. nov., B flavifrons group A. oralis (Navás, 1914) comb. nov., B ? A. parabola (Okamoto, 1919) comb. nov., B alcestes group A. perfecta (Banks, 1895) comb. nov., B flavifrons group A. perpallida (Tjeder, 1966) comb. nov., B flavifrons group A. pervenosa (Tjeder, 1966) comb. nov., B ? primary type a female A. phantosula (X.-k. Yang & C.-k. comb. nov., B ? Yang, 1990) ...Continued on the next page

222 · Zootaxa 4966 (2) © 2021 Magnolia Press BREITKREUZ et al. Table 1. (Continued) Species Author(s), Year Status, Species Group Notes Category A. phlebia (Navás, 1927) comb. nov., B venosa group* A. physophlebia (Navás, 1914) A ? may not belong in Apertochrysa

A. picteti (McLachlan, 1880) comb. nov., B flavifrons group A. pieli (Navás, 1931) comb. nov., B ? A. pilinota (Dong et al., 2004) comb. nov., B ? A. prasina (Burmeister, 1839) comb. nov., B prasina group A. pulchrina (Tjeder, 1966) comb. nov., B alcestes group A. puncticollis (Banks, 1940) A ? tignum absent A. punctilabris (McLachlan, 1894) comb. nov., B ? A. punctithorax (New, 1980) A ? may not belong in Apertochrysa

A. qingchengshana (C.-k. Yang et al., comb. nov., B ? 1992) A. qinlingensis (C.-k. Yang & X.-k. comb. nov., B ? Yang, 1989) A. raedarii (Hölzel & Ohm, comb. nov., B flavifrons group* 2000) A. rothschildi (Navás, 1915) comb. nov., B ? A. rubicunda (Hölzel, 1993) comb. nov., B venosa group A. rubra (Hölzel et al., 1994) comb. nov., B ? primary type a female A. sana (X.-k. Yang & C.-k. comb. nov., B ? Yang, 1990) A. sansibarica (Kolbe, 1897) comb. nov., B ? A. selenia (Navás, 1912) comb. nov., B flavifons group*? A. sensitiva (Tjeder, 1940) comb. nov., B venosa group* A. setosa (Hölzel & Ohm, comb. nov., B ? primary type a female 1995) A. sierra (Banks, 1924) comb. nov., B flavifrons group* A. spadix (Hölzel, 1988) comb. nov., B venosa group* A. spissinervis (Tjeder, 1966) comb. nov., B flavifrons group A. subcostalis (McLachlan, 1882) comb. nov., B venosa group* A. subcubitalis (Navás, 1901) comb. nov., B flavifrons group* A. subflavifrons (Tjeder, 1949) comb. nov., B venosa group* A. sybaritica (McLachlan in comb. nov., B venosa group*? Fedchenko, 1875) A. tacta (Navás, 1921) comb. nov., B venosa group A. teiresias (Hölzel & Ohm, comb. nov., B flavifrons group? 1982) A. triactinata (New, 1980) A — correct genus uncertain

A. triangularis (Adams, 1978) comb. nov., B ? ...Continued on the next page

Apertochrysa, a new senior synonym of Pseudomallada Zootaxa 4966 (2) © 2021 Magnolia Press · 223 Table 1. (Continued) Species Author(s), Year Status, Species Group Notes Category A. triangularis (C.-k. Yang & X.-x. comb. nov., B ? an unreplaced junior secondary Wang, 1994) homonym of triangularis (Adams) in both Pseudomallada and Apertochrysa A. tridentata (X.-k. Yang & C.-k. comb. nov., B alcestes group* Yang, 1990) A. truncatata (X.-k. Yang et al. comb. nov., B ? in X.-k. Yang et al., 2005) A. umbrosa (Navás, 1914) A venosa group tignum absent; type species of Apertochrysa A. ussuriensis (Makarkin, 1985) comb. nov., B alcestes group A. varians (Kimmins, 1959) comb. nov., B ? A. venosa (Rambur, 1838) comb. nov., B venosa group A. venosella (Esben-Petersen, comb. nov., B venosa group* 1920) A. ventralis (Curtis, 1834) stat. nov., C prasina group A. venusta (Hölzel, 1974) comb. nov., B venosa group* A. verna (C.-k. Yang & X.-k. comb. nov., B prasina group* Yang, 1989) A. viridifrons (Hölzel & Ohm, comb. nov., B venosa group* 1999) A. vitticlypea (C.-k. Yang & X.-x. comb. nov., B ? Wang, 1990) A. waitei (Tillyard, 1917) A — Mallada sp.? A. wangi (C.-k. Yang et al., comb. nov., B ? 1992) A. wuchangana (C.-k. Yang & X.-x. comb. nov., B ? Wang, 1990) A. xiamenana (C.-k. Yang et al., comb. nov., B ? 1999) A. yangi (X.-k. Yang & X.-x. comb. nov., B ? Wang in X.-k. Yang et al., 2005) A. yunnana (C.-k. Yang & X.-x. comb. nov., B alcestes group* Wang, 1994) A. yuxianensis (Bian & Li, 1992) comb. nov., B alcestes group* A. zelleri (Schneider, 1851) comb. nov., B prasina group A. zulu (Tjeder, 1966) comb. nov., B ? primary type a female

References

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